King Crabs of New Zealand, Australia and the Ross Sea (Crustacea: Decapoda: Lithodidae)

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The Marine Fauna of New Zealand: King Crabs Zealand, Australia and ISSN 2463-638X; 123 (Online) the Ross Sea (Crustacea: Decapoda: Lithodidae)

The Marine Fauna of New Zealand: Shane T. Ahyong King Crabs of New Zealand, Australia and the Ross Sea (Crustacea: Decapoda: Lithodidae)

Shane T. Ahyong Soft cover: ISBN 978–0–478–23285–1 Hard cover: ISBN 978–0–478–23286–8 NIWA Biodiversity Memoir 123

The Marine Fauna of New Zealand: King Crabs of New Zealand, Australia and the Ross Sea (Crustacea: Decapoda: Lithodidae)

Shane T. Ahyong

Marine Biodiversity and Biosecurity National Institute of Water and Atmospheric Research Ltd Private Bag 14901, Kilbirnie Wellington, New Zealand

NIWA Biodiversity Memoir 123

2010

AHYONG, S. T. The marine fauna of New Zealand: King crabs of New Zealand, Australia and the Ross Sea (Crustacea: Decapoda: Lithodidae) / by Shane T. Ahyong—Wellington: NIWA (National Institute of Water and Atmospheric Research Ltd), 2010 (NIWA Biodiversity Memoir, ISSN 1174-0043; 123)

Soft cover: ISBN 978–0–478–23285–1 Hard cover: ISBN 978–0–478–23286–8

Series Editor: Dennis P. Gordon Copy edited by: Tracy Farr Typeset and indexed by: Geoff Gregory, Word Therapy, Paraparaumu Printed and bound by: Graphic Press & Packaging Ltd, Levin

Received for publication—15 February 2010

The Marine Fauna of New Zealand: King Crabs of New Zealand, Australia and the Ross Sea (Crustacea: Decapoda: Lithodidae)

Shane T. Ahyong

Marine Biodiversity and Biosecurity National Institute of Water and Atmospheric Research Ltd Private Bag 14901, Kilbirnie Wellington, New Zealand

ABSTRACT

The king crabs, Lithodidae, are among the largest and most conspicuous deepwater crustaceans worldwide, including the polar regions. Prior to the present study, 10 extant species were believed to occur in New Zealand and Australia: Lithodes murrayi, Lithodes longispina, Neolithodes brodiei, Neolithodes flindersi, and six species of Paralomis. Three species in two genera of lithodid have been recorded from the Ross Sea, Antarctica. Surprisingly, however, the Australian and New Zealand lithodid fauna has never been studied in detail and the identities of most species in the region have never been explicitly scrutinised. The Lithodidae from New Zealand, Australia, and the Ross Sea are thus fully reviewed. A total of three genera and 23 species are now known from the study area: 13 species in New Zealand, 12 in Australia (four shared by both regions) and three in the Ross Sea (of which one species extends onto the Macquarie Ridge). Neither L. murrayi nor L. longispina occurs in the region. Previous records of L. longispina and L. murrayi from New Zealand are based on different ontogenetic phases of the same undescribed species, herein named L. aotearoa sp. nov. Previous records of L. longispina from Australia are based on Lithodes richeri and an undescribed species, herein named L. australiensis sp. nov. Records of Lithodes murrayi from Macquarie Island are referable to L. macquariae sp. nov. Thus, L. longispina is reliably known only from Japan, and L. murrayi is reliably known only from the southwestern Indian Ocean. Lithodes murrayi does not exbihit a circumpolar distribution as formerly believed. Neolithodes brodiei is known only from New Zealand; reports of the species from Australia are based on the recently described N. flindersi Ahyong, 2010. Previous reports of Paralomis histrix from New Zealand are based on juvenile Paralomis zealandica, and P. cf. histrix from southeastern Australia are referable to a new species, P. echidna sp. nov. In total 14 new species are described, five exclusively from New Zealand, five exclusively from Australia, and four shared by both regions. Of the three lithodid species known from the Ross Sea, Neolithodes yaldwyni Ahyong & Dawson, 2006 and Paralomis stevensi Ahyong & Dawson, 2006 are presently known only from Antarctic waters; the third species, P. birsteini Macpherson, 1988 ranges onto the Macquarie Ridge. Paralomis stevensi is reported for the first time from the Amundsen Sea.Paralomis shinkaimaruae Takeda & Hatanaka, 1984 is synonymised with P. zealandica Dawson & Yaldwyn, 1971. Results of the present study demonstrate considerably higher species richness than previously known, essentially doubling the known fauna of the region.

Keywords: Crustacea, Decapoda, Lithodoidea, Lithodidae, king crabs, systematics, taxonomy, new species, Australia, New Zealand, Antarctica

Abstract...... 3 Introduction...... 5 Materials and Methods...... 7 Morphology...... 8 Systematics...... 14 Lithodoidea Samouelle, 1819...... 14 Key to families of Lithodoidea Samouelle, 1819...... 14 Lithodidae Samouelle, 1819...... 14 Key to genera of Lithodidae Samouelle, 1819...... 14 Lithodes Latreille, 1806...... 15 Key to adult Lithodes from New Zealand and Australia ...... 16 Lithodes aotearoa sp. nov...... 16 Lithodes australiensis sp. nov...... 29 Lithodes chaddertoni sp. nov...... 37 Lithodes jessica sp. nov...... 41 Lithodes macquariae sp. nov...... 48 Lithodes rachelae sp. nov...... 55 Lithodes richeri Macpherson, 1990...... 60 Lithodes robertsoni sp. nov...... 66 Neolithodes A. Milne-Edwards & Bouvier, 1894...... 73 Key to species of Neolithodes from New Zealand, Australia and the Ross Sea...... 73 Neolithodes brodiei Dawson & Yaldwyn, 1970...... 74 Neolithodes bronwynae sp. nov...... 83 Neolithodes flindersi Ahyong, 2010...... 89 Neolithodes yaldwyni Ahyong & Dawson, 2006...... 101 Paralomis White, 1856...... 108 Key to adult Paralomis from New Zealand, Australia and the Ross Sea...... 109 Paralomis birsteini Macpherson, 1988...... 110 Paralomis dawsoni Macpherson, 2001...... 116 Paralomis echidna sp. nov...... 125 Paralomis gowlettholmes sp. nov...... 135 Paralomis hirtella de Saint Laurent & Macpherson, 1997...... 142 Paralomis poorei sp. nov...... 148 Paralomis staplesi sp. nov...... 156 Paralomis stevensi Ahyong & Dawson, 2006...... 160 Paralomis taylorae sp. nov...... 166 Paralomis webberi sp. nov...... 169 Paralomis zealandica Dawson & Yaldwyn, 1971...... 175 Discussion...... 185 Acknowledgments...... 186 References...... 187 species INDEX...... 193

The king crabs, Lithodoidea, comprise a small group (McLaughlin & Lemaitre 1997; McLaughlin et al. 2004) (~100 species) of largely deepwater species, of which and proposed a distant relationship between lithodids more than 60 occur in the Pacific Ocean. They are best and other paguroids (McLaughlin et al. 2007). In con- known on the basis of several commercially important trast, most other phylogenetic studies have recovered northern hemisphere species, especially Paralithodes a close relationship between lithodids and pagurids camtschaticus (Tilesius, 1815) (Red King Crab) and Lith- supporting the derivation of king crabs from shell- odes aequispinus (Benedict, 1895) (Golden King Crab). dwelling hermit crabs (Richter & Scholtz 1994; Mor- In the southern hemisphere, Lithodes santolla (Molina, rison et al. 2002; Ahyong & O’Meally 2004; Tsang et al. 1782) (Southern King Crab) and several species of 2008; Ahyong et al. 2009; Bracken et al. 2009). As such, Paralomis are commercially landed off southern South recognising the superfamily Lithodoidea, distinct from America, and exploratory fisheries for lithodids are Paguroidea, as proposed by McLaughlin et al. (2007) currently being undertaken in New Zealand waters. is not compatible with the weight of phylogenetic Although the lithodoids are exceptionally crab-like in evidence gathered to date. However, pending ongo- appearance, they are not true crabs (infraorder Brachy- ing studies of anomuran phylogeny, the classification ura), rather being part of the infraorder Anomura, of McLaughlin et al. (2007) is followed here, with which includes squat lobsters and hermit crabs. Until Lithodoidea treated as a superfamily, and Lithodidae 2007, the king crabs comprised the family Lithodi- restricted to non-hapalogastrine king crabs. Irrespec- dae, containing two subfamilies, Lithodinae and tive of the phylogenetic position of the king crabs, all Hapalogastrinae, within the superfamily Paguroidea. workers recognise the validity of the two major clades, McLaughlin et al. (2007), however, regarded the king corresponding to Lithodidae and Hapalogastridae, crabs as only distant relatives of the hermit crabs, and respectively. raised the Lithodidae and its constituent subfamilies to superfamilial and familial rank, respectively — Lithod- World studies oidea, containing Lithodidae and Hapalogastridae. Hapalogastrids occur only in the northern Pacific, so Prior to the last three decades, most research into the the present study deals only with Lithodidae sensu Lithodidae was focused on the northern hemisphere McLaughlin et al. (2007). fauna. Key studies include Bouvier (1895, 1896), The lithodoids are the paradigm example of carcin- Benedict (1895), Hansen (1908), Schmitt (1921), and isation, that is, the evolution of a crab-like form from a Makarov (1938). Enrique Macpherson (1988b) revised non-crab-like ancestor. No other decapods more closely the Atlantic fauna, covering 30 species, and studied approximate the brachyuran form than do lithodoids. smaller collections from the Indian Ocean and south- The process of attaining crab-like form is typically western Pacific Ocean (e.g. Macpherson 1988a, c, 1989, understood to involve a shortening and broadening 2001, 2003; Macpherson & Chan 2008). Lithodids from of the carapace (from an elongate, often subcylindri- Antarctic and sub-Antarctic waters have been de- cal form), and a reduction of the abdomen, which is scribed by Macpherson (2004), Spiridonov et al. (2006), held folded beneath rather than extended behind the and Ahyong & Dawson (2006). Most recently, lithodids cephalothorax. Lithodoids have long been thought have been studied by Takeda & Nagai (2004), Takeda to be derived from within the asymmetrical hermit & Bussarawit (2007), Hall & Thatje (2009b), Ahyong & crabs, the so-called ‘hermit to king’ hypothesis (Bou- Chan (2010), Ahyong (2010) and Ahyong et al. (2010) vier 1894a, b, 1897; Cunningham et al. 1992). Indeed, from various Atlantic and Indo-Pacific localities. the close relationship of lithodids to asymmetrical Dawson (1989) provided a useful world bibliography hermit crabs, particularly to pagurids, has long been of the references dealing with Lithodidae, and Zaklan widely accepted through the placement of lithodoids (2002) provided a synoptic overview of the biology and among the paguroids in most classification schemes taxonomy of the group. of the Anomura (Martin & Davis 2001). The primary evidence used for this argument was the asymmetry Previous regional studies of the somites and pleopods found in the female abdomen of lithodoids. Several recent studies, however, Studies of Antipodean decapods date back more have challenged the ‘hermit to king’ hypothesis, cit- than a century (e.g. Heller 1865; Hess 1865; Miers ing apparently implausible transformation pathways 1876; Haswell 1882; Hutton 1882; Filhol 1883, 1885a),

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ though lithodids were discovered in New Zealand P. cf. phrixa and Lithodes aff. longispina from southwest- and Australian waters only much more recently. The ern Australia. Including the record of L. murrayi from first lithodid to be recorded in the region, identified Macquarie Island, six species have been reported from as Lithodes murrayi Henderson, 1888 by Hale (1941), Australian waters, of which three were apparently was collected in 1930 near Macquarie Island (south shared with New Zealand (Poore 2004). Ahyong (2010) of Tasmania) by the British, Australian and New Zea- showed that Australian records of N. brodiei were based land Antarctic Research Expedition (BANZARE). No on an undescribed species, named N. flindersi Ahyong, other lithodids were recorded from the region until 2010. Otherwise, no further progress on the taxonomy 1958, when a juvenile specimen of Paralomis zealan- of Lithodidae in New Zealand and Australian waters dica from Cook Strait, New Zealand, identified then has been made, although significant collections in both as Lithodes sp., was figured by King (1958). In 1960 a countries have been accumulated. specimen, identified asLithodes murrayi, was captured Three species of lithodids are known from the Ross by crayfishers in Foveaux Strait, New Zealand (Dell Sea region, Antarctica (Ahyong & Dawson 2006). The 1963; Yaldwyn & Dawson 1970). Since then, increased first lithodid recorded from the Ross Sea (near Scott deepwater exploration in New Zealand waters by the Island) was identified as the North Atlantic Paralomis New Zealand Oceanographic Institute (now NIWA) spectabilis Hansen, 1908 (Birstein & Vinogradov 1967), led to the discovery of not insignificant populations an apparently bi-polar species. Macpherson (1988a), of lithodids, among them Neolithodes brodiei Dawson however, showed that the southern records of P. & Yaldwyn, 1970 and Paralomis zealandica Dawson & spectabilis were referable to a distinct species, Paralomis Yaldwyn, 1971. Subsequently, other species were also birsteini Macpherson, 1988a. More intensive sampling reported from New Zealand: Lithodes longispina Sakai, of the Ross Sea over the last half decade by NIWA 1971, Paralomis histrix (de Haan, 1844), Paralomis hirtella and the New Zealand Ministry of Fisheries recovered de Saint Laurent & Macpherson, 1997, and Paralomis two further species, the known lithodid fauna now dawsoni Macpherson, 2001 (see Dawson & Yaldwyn comprising Paralomis birsteini Macpherson, 1988a, 1985; Webber & Naylor 2004a, b; Ahyong et al. 2007; P. stevensi Ahyong & Dawson, 2006, and Neolithodes Dawson 2008). Additionally, the first and only known yaldwyni Ahyong & Dawson, 2006. Note that Davie fossil lithodid was described from mid-late Miocene (2002) inadvertently listed Lithodes murrayi from the formations of New Zealand, Paralomis debodeorum Feld- Possession Island in the Ross Sea, rather than Posses- mann, 1998. In total, seven names have been applied to sion in the southwestern Indian Ocean. extant species of Lithodidae in New Zealand. Thus, a total of 13 nominal lithodid species have In Australia, lithodids were not known from been reported from the study area, although only the mainland waters until the 1980s when deepwater ex- Ross Sea fauna has been well characterised. This study ploration off New South Wales (by New South Wales commenced as a revision of the New Zealand Lithodi- Fisheries) and off Victoria and Tasmania (by the Com- dae. However, the presumed faunal overlap between monwealth Scientific and Industrial Research Organi- New Zealand, Australian and Antarctic species, along sation, CSIRO) discovered five species of lithodids on with taxonomic ambiguity of several nominal species, seamounts, outer shelf and slope waters. These species required revision of the regional fauna. Thus, the extant were reported as Lithodes longispina, Neolithodes brodiei, Lithodidae of New Zealand, Australia and the Ross Sea Paralomis cf. birsteini, P. cf. histrix, and P. cf. phrixa (see are revised below. Poore 2004). Poore et al. (2008) subsequently recorded

MATERIALS AND METHODS

Study area and study material by numerous commercial fishing vessels. For economy, the vessel name is not explicitly mentioned under Ma- The study area encompasses the region bounded by terial Examined; Tangaroa and Kaharoa station numbers New Zealand Exclusive Economic Zone (EEZ), south- are prefaced by TAN and KAH respectively. In other ern Australia (south of 30°S), including the Macquarie instances, the vessel name is mentioned. Lithodids col- Ridge, and the Ross Sea, Antarctica (Fig. 1). Lithodid lected from Australian waters were collected by New collections held in the National Institute of Water and South Wales Fisheries (FRV Kapala) along the southeast Atmospheric Research, Wellington, New Zealand coast, various CSIRO research programmes (RV Soela, (NIWA), Museum of New Zealand Te Papa Tongarewa RV Southern Surveyor), especially between Tasmania (NMNZ), Australian Museum (AM), Institut de recher- and Western Australia, and miscellaneous commercial che pour le développement, Nouméa (IRD), Museum fishing vessels. All images are by the author unless Victoria (NMV), Tasmanian Museum (TM), and South indicated otherwise. Australian Museum (SAM) were comprehensively studied, along with type and other specimens in the col- Measurements and terminology lections of the Muséum national d’Histoire naturelle, Paris (MNHN), National Science Museum, Tokyo Morphological terminology is discussed below and il- (NSMT), National Taiwan Ocean University, Keelung lustrated in Figs 2–4. Carapace length (cl) is measured (NTOU), Raffles Museum of Biodiversity Research, along the dorsal midline from the apex of the rostrum National University of Singapore (ZRC), National to the posterior carapace midline. Postorbital carapace Museum of Natural History, Smithsonian Institution, length (pcl) is measured from the posterior margin of Washington D.C. (USNM), Florida Museum of Natural the orbit to the posterior margin of the carapace. Cara- History (FLMNH), and Zoological Laboratory of the pace width (cw) is greatest carapace width excluding Kyushu University, Fukuoka, Japan (ZLKU). The ma- spines. Other measurements are shown in Fig. 4. All jority of specimens from the New Zealand region were measurements are in millimetres (mm). Distribution collected by various benthic surveys conducted by maps were produced with the aid of Online Map Crea- NIWA (formerly NZOI) (RV Tangaroa, RV Kaharoa) and tion (www.aquarius.ifm-geomar.de).

Characters currently used in lithodid taxonomy are de- anterolateral spine; and posteriorly, a long spine adja- rived from the cephalothorax, especially the antennae cent to the junction with the branchial margin, termed and ornamentation of the carapace; the ornamentation the hepatic spine. At any given body size, the hepatic and proportions of the chelipeds (pereopod 1) and spine is usually the longest of the carapace spines. The walking legs (pereopods 2–4); and abdomen. Owing branchial margin typically bears several major spines to the considerable allometric changes that typically (longer than adjacent spines, especially in juveniles) occur between juvenile and adult stages, diagnoses that serve as useful landmarks: one above pereopod 3 and descriptions given under taxonomic accounts are (primary marginal branchial spine 1), and one above based on adults. Juvenile morphology is discussed in pereopod 4 (primary marginal branchial spine 2). the Remarks for each species. These large spines effectively divide the branchial margin into discrete regions, herein termed the ante- Cephalothorax: The cephalothorax comprises an rior branchial margin, lateral branchial margin, and acron (bearing the eyes), five cephalic somites bearing posterior branchial margin. The two branchial regions appendages (antennule, antenna, mandible, maxil- are separated anteriorly by the gastric and cardiac re- lule, and maxilla) and eight thoracic somites bearing gions, and posteriorly by the intestinal region, marked maxillipeds 1–3 and pereopods 1–5. These somites by paired intestinal spines in Lithodes and Neolithodes. are fused and entirely covered by the carapace. The The margin between each of these major spines is lined cephalothorax is ventrally represented by a series of with varying numbers of shorter spines. The number fused sternal plates. In most lithodids, the cephalot- of posterior branchial marginal spines is diagnostic in horax is covered by a well-calcified carapace which species of Lithodes. is generally pentagonal or pyriform. A prominent The dorsal surface of the carapace is divided into rostrum is present anteromedially. In Paralomis and regions by shallow grooves. The most obvious is the Neolithodes, the rostrum is trifid, consisting of a median cervical groove, which crosses the carapace and sepa- spine and a pair of divergent dorsal spines. In Lithodes, rates the gastric and hepatic regions from the branchial the rostral structure differs from that of Paralomis and and cardiac regions. The gastric region is higher than Neolithodes, having a strong anterior projection with a the other regions and lies behind the rostrum; it usually pair of dorsal spines placed at some distance from the bears four major spines (most obvious in Lithodes and base, demarcating an anterior and posterior portion. Neolithodes but not always obvious in other genera) The anterior portion is itself distally bifid, though the in addition to other spinules or granules that may be distal spines are sometimes absent owing to damage present. The regions flanking the gastric region are or moulting abnormalities. The posterior portion also termed hepatic. The cardiac region lies medially behind bears a prominent ventral, anteriorly directed spine, the gastric region and bears two or four major spines. herein termed the ventral rostral spine. The ventral ros- The cardiac region is followed by the intestinal region tral spine in Lithodes appears to be homologous with the and usually bears one or two spines along the midline. median rostral spine of Paralomis and Neolithodes, but The intestinal region is often ill-defined and difficult different terminology is employed here for taxonomic to distinguish from the branchial or cardiac regions. convenience. In most species of Lithodes, the ventral The branchial regions flank the cardiac and intestinal rostral spine is not visible dorsally, but in L. santolla regions and typically bear three major spines of which (Molina, 1782) and L. confundens Macpherson, 1988b the anterior two are longer. The spine pattern is con- the anterior and posterior dorsal portions are much sistent across most lithodids, and homologous spines reduced in length, and are distally overreached by the can be identified across species and genera, except in ventral rostral spine. As a result, in dorsal view, the species in which the carapace is entirely and densely anterior dorsal portion may appear to be distally trifid, covered with spines or tubercles, or dorsally unarmed rather than bifid as it actually is. as in Paralomis zealandica, P. hirtella, and P. dawsoni The orbital margin is delimited medially by the ros- (although spines may be present in early juveniles). trum and laterally by the outer orbital spine. The eyes As is well known in lithodids, the length of carapace have a unisegmental peduncle (ocular peduncle) and spines changes dramatically between juveniles and well-pigmented, distoventrally positioned cornea. The adults, and must be considered during the identifica- ocular peduncle usually bears small spines or granules. tion process. As indicated by Macpherson (1988b), The hepatic region bears, at the anterolateral angle, the however, although the proportional length of spines

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ varies considerably with size, the number and position In the Hapalogastridae (not known from the study of spines is remarkably constant within most species. area), the tergites of somites 3–5 are entirely uncalcified. Thoracic sternum: The visible thoracic sternites are represented ventrally by the fused sternites 4–7 form- Antennule: The antennule is the first cephalic ap- ing the sternal plastron. Sternal characters are not pendage, and consists of a 3-segmented peduncle and used in lithodoid taxonomy, although the presence of paired flagella. The basal article bears the statocyst. a deep median fissure on sternite 5 (between the first The distal two articles are typically cylindrical and pair of walking legs) is useful at generic level, and will unarmed. Two multi-segmented flagella arise api- separate Lithodes and Neolithodes (fissure present) from cally from the ultimate peduncular article. The upper Paralomis (fissure absent). flagellum is long and bears numerous long aesthetascs along its ventral margin. The lower flagellum is short Abdomen: The abdomen is short, broad, variously and small-segmented. calcified, and folded underneath the cephalothorax. It consists of six somites plus telson. The terga are vari- Antenna: The antenna is the second cephalic append- ously calcified but the sternal surfaces are membranous age and consists of a 5-segmented peduncle and long, and uncalcified. Males lack pleopods, but females often uniramous flagellum. The proximal articles of the have a pair of small first pleopods and uniramous, peduncle are usually spinose. The first article bears unpaired left pleopods 2–5. Uropods are absent. the urinal or green gland, opening anteromesially. Abdominal somite 1 is reduced and often armed The exopod, termed the scaphocerite, is present in with a pair of submedian spines. Somite 2 is widest most lithodids including Paralomis, but reduced to a and deepest, and bears important diagnostic features. small sclerite or spine in Lithodes and Neolithodes. The It is plesiomorphically divided into five plates: median, scaphocerite, when present, varies from being a simple paired submedian, and paired marginals. In Neolith- spine with a few subsidiary spinules, to an elaborate odes, all five plates are separate and demarcated by a multispinous structure. suture. The median and submedian plates are large and broad whereas the marginal plates are very narrow. In Pereopod 1 (chelipeds): The first pereopods are the Paralomis, all five plates are indistinguishably fused chelipeds and usually asymmetrical in size, especially into a single large plate. In Lithodes, the median and in males. In females, the size difference in the chelipeds submedian plates are always indistinguishably fused is much less pronounced. The right cheliped is the into a single plate (the ‘central’ plate), but the marginal larger (rarely vice versa), as in Paguridae. The cheliped plates may or may not be fused with the central plate. consists of dactylus, propodus, carpus, merus, ischio- Where the marginal plates are fused with the remain- basis, and coxa. Unlike the walking legs (pereopods ing plates, a shallow crease is usually visible indicating 2–4) in which the segments more-or-less articulate in the line of fusion. The condition of the marginal plate the same plane, the arrangement of the cheliped seg- of somite 2 (fused or unfused) is a useful diagnostic ments allow articulation in multiple planes, making character for adults. The marginal plates are always positional terminology such as extensor and flexor free in juveniles, even in species in which the plates margins difficult to apply. Therefore, in the case of the are fused as adults (e.g. Lithodes aotearoa). Tergites 3–5 chelipeds, terminology used here refers to the position are symmetrical in males, and asymmetrical in females of structures when the cheliped is held forwards with (via enlargement of the left plates and reduction of the dactylus uppermost. the right plates), but are variously calcified between lithodid genera. In Paralomis, the terga are composed Pereopods 2–4 (walking legs 1–3): Pereopods 2–4 are of median, submedian, and marginal plates, of which the walking legs, and their spination, relative overall the marginals are often subdivided (females with left lengths and the relative segment proportions are often marginals fused with submedians). In Lithodes, the diagnostic at species level. The walking legs are simi- median plates are composed of blunt nodules, the sub- lar in structure and ornamentation, differing slightly medians are entire, and the marginals are subdivided in length and proportions. They consist of dactylus, (females with left marginals fused with submedians). propodus, carpus, merus, ischiobasis, and coxa. The The terga in Neolithodes are membranous and densely length of the walking legs is sexually dimorphic, with covered with spinular or nodular calcified elements; in that of males usually about 10–15% longer than in size- females, the submedians and marginals are fused into matched females. The lengths and proportions of walk- a single plate. Tergite 6 comprises an elongate central ing leg segments, especially the merus and propodus, plate and narrow, subdivided marginal plates. The vary allometrically, typically being more slender in telson is usually short and semicircular. juveniles than adults. As in other Anomura, the female

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 2. General morphology. A, habitus. B–C, carapace spines and regions. D, rostrum features of Paralomis and Neolithodes. E, antenna.

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 3. Thoracic and abdominal morphology. A, ventral surface of thoracic sternum and pereopodal coxae. B, C, D, abdominal features of Lithodes, Neolithodes, and Paralomis, repectively; from left to right: abdominal somite 2 in posterior view, male abdominal somites 2–6 and telson (T), female abdominal somites 3–6 and telson. 11

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 4. Pereopodal morphology and carapace measurements. A–B, cheliped (pereopod 1) morphology and measurements. C–D, walking leg (pereopods 2–4) morphology and measurements. E, carapace measurements.

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ gonopore is located on the pereopod 3 coxa, and male of the walking leg dactyli. The dactyl length relative gonopore on the pereopod 5 coxa. Mature females to propodus length decreases with increasing body usually have a setose surface of the pereopodal coxae. size, and in juveniles, the flexor margins are lined with In juveniles, especially of Lithodes and Neolithodes, the a row of corneous spines. These corneous spines are dactyli of the walking legs are proportionally longer retained in adult Paralomis, but lost in early juvenile (in comparison to the propodus) in juveniles than in Neolithodes and adults of most species of Lithodes (be- adults. The flexor margins of the walking leg dactyli coming diagnostic for those species retaining the spines in juveniles through adults bear a row of corneous as adults, e.g. L. chaddertoni sp. nov. and L. jessica sp. spinules in Paralomis, and are unarmed in early juve- nov.). In most species of Lithodes, the marginal plates of niles through to adults in Neolithodes. Juvenile Lithodes abdominal somite 2 are separated from the submedian have a row of corneous spinules on the flexor margins, plates by a suture in juveniles that is present through which, in most species, is lost in late juveniles. The to adults. In adults of some species, however, such as effects of allometry and sexual dimorphism must be L. aotearoa sp. nov. and L. richeri Macpherson, 1990, the carefully considered when identifying specimens. marginals are fused to the submedians. Thus, fusion or separation of the plates of abdominal somite 2 can Pereopod 5 (eighth thoracic appendage): Pereopod 5 be a useful diagnostic feature in adults. is a very small and slender subchelate appendage, and Sexual dimorphism is evident but less pronounced is carried folded underneath the carapace. It is used than allometric changes. Primary sexual differences, for grooming the gills and the eggs in females, and, at such as abdominal symmetry or asymmetry and present, has no taxonomic value. gonopore position, as discussed above, are the most obvious differences. However, secondary sexual dif- Allometry and sexual dimorphism ferences are present in the form of pereopod length and shape, and these differences become increasingly When identifying lithodids, particular attention must pronounced with size. The chelipeds of males are be given to the size and sex of specimens (see Figs usually distinctly unequal (except in Lithodes jessica), 2–4 for morphological terms and measurements used with the right chela larger and more robust than the herein). Lithodids exhibit considerable allometric vari- left chela. In females, both chelae are similar, having ation in rostral and general spine length. These features the form of the left chela of males. The walking legs nevertheless provide very useful diagnostic characters, differ in length between sexes, being proportionally provided body size is considered. The rostrum and longer in males than in females. Typically, they differ general spines are proportionally longest in juveniles, by about 10–15%, sometimes more, being most obvious becoming progressively shorter proportionally (and in mature, size-matched specimens. Another factor to sometimes absolutely) with increasing body size. These consider when identifying specimens is the the effect allometric changes are most pronounced in species of parasitic feminisation caused by rhizocephalan of Lithodes and Neolithodes, in which the rostrum and infections. The abdomen of specimens should always longest carapace spines may exceed the postorbital be checked for the presence of rhizocephalan externae. carapace length (pcl) in early juveniles. Allometric In males with advanced rhizocephalan infection, the trends in Paralomis are similar to, but less pronounced pereopod morphometrics are similar to females. than in Lithodes and Neolithodes. In addition to reduc- To best facilitate identification of species, figures of tion in relative spine length, a number of allometric both sexes, across their known size range, are provided changes are also evident in length and ornamentation where possible.

LITHODOIDEA Samouelle, 1819 Composition. Cryptolithodes Brandt, 1848, Glyptolith- odes Faxon, 1895, Lithodes Latreille, 1806, Lopholithodes Lithodiadae Samouelle, 1819: 90. Brandt, 1848, Neolithodes A. Milne-Edwards & Bouvier, Lithodoidea. — McLaughlin et al., 2007: 108. 1894b, Paralithodes Brandt, 1848, Paralomis White, 1856, Phyllolithodes Brandt, 1848, Rhinolithodes Brandt, 1848, Diagnosis. Crab-like. Carapace subtriangular to trans- Sculptolithodes Makarov, 1934. versely ovate; rostrum present. Eyes with pigmented cornea; cornea distoventral on stalk. Maxilliped 3 Remarks. Lithodidae includes 10 genera. The three with crista dentata and accessory tooth. Pereopods largest genera, Lithodes, Neolithodes, and Paralomis, are 1 chelate, subequal or right larger and more robust represented in New Zealand, Australian and Antarctic than left. Pereopods 2–4 developed as walking legs. waters. Phylogenetic relationships between lithodid Pereopod 5 subchelate, reduced, usually folded within genera were most recently studied by Hall & Thatje branchial chamber. Abdomen short, broad, held folded (2009a), who found Neolithodes and Lithodes to be more against thoracic sternum; composed of calcified plates closely related to each other than either is to Paralomis. or nodules embedded in arthrodial membrane; somite The genera of Lithodidae are distinguished in the key 1 largely concealed by posterior carapace margin; ab- below, with those represented in New Zealand marked dominal plates more strongly developed on left side in bold. in females, producing asymmetry. Pleopods absent in males; females with paired pleopods on somite 1 and Key to genera of Lithodidae single uniramous pleopod on left side of somites 2–5. Uropods absent. 1. Carapace nearly smooth, without protuberances, granules, or spines; markedly wider than long, Composition. Lithodidae Samouelle, 1819, Hapal- expanded laterally covering walking legs entirely ogastridae Brandt, 1850. when the legs are drawn in. Rostrum broad, compressed, truncated anteriorly...... Cryptolithodes Key to families of Lithodoidea Samouelle, 1819 – Carapace with more or less numerous protuber- 1. Rostrum well-developed, spiniform or truncate, ances, granules or spines. Lateral margins of cara- extending anteriorly beyond eye. Abdominal pace without lateral expansions overhanging legs. somites 3–5 usually well-calcified; sometimes with Walking legs never completely covered by carapace membranous areas medially with nodules or small when drawn in close to body. Rostrum variable in spinules...... Lithodidae shape, but never truncated anteriorly...... 2 – Rostrum short, broad, not usually extending ante- 2. Sternite 5 (sternite of first pair of walking legs) with riorly beyond eye. Abdominal somites 3–5 ill-calci- a deep, longitudinal, medial fissure...... 3 fied, in form of a membranous sac; usually covered – Sternite 5 (sternite of first pair of walking legs) by small, thin plates...... Hapalogastridae without a deep, longitudinal, medial fissure...... 5 3. Abdominal somite 2 composed of three plates LITHODIDAE Samouelle, 1819 (paired marginal, central plate formed by fused median and submedians) or fused into a single Lithodiadae Samouelle, 1819: 90 [type genus: Lithodes La- plate...... Lithodes treille, 1806] [incorrect original spelling for Lithodidae Samouelle, 1819 (ICNZ Opinion 511)] – Abdominal somite 2 composed of five plates (me- Lithodinae. — Macpherson, 1988b: 17. dian, paired submedian, and paired marginal.....4 Lithodidae. — McLaughlin et al., 2007: 108. — De Grave et al., 2009: 25. 4. Abdominal somites 3–5 comprising small, more or less spiniform nodules in males, connected by Diagnosis. Rostrum well-developed, spiniform or arthrodial membrane, without well-defined plates. truncate, extending anteriorly beyond eye. Abdominal Females with a well-developed submedian plate somites 3–5 usually well-calcified; composed entirely on the left side of each segment, right side either of calcified plates or of nodules or spinules embedded with spiniform nodules connected by arthrodial in arthrodial membrane. membrane or with plates fused. Scaphocerite a

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ small spine or reduced to a short sclerite...... present. Abdominal somite 2 comprising 3 plates (a ...... Neolithodes wide ‘central’ plate and a pair of narrow marginal plates) or single plate in adults of some species (all – Abdominal somites 3–5 bearing calcified nodules plates fused). Abdominal somites 3–5 with median medially, with well-defined submedian and mar- regions composed of calcareous nodules joined by ginal plates on each side in both sexes. Scaphocerite arthrodial membrane; submedian and marginal plates well-developed, spinous...... Paralithodes well demarcated in both sexes; submedian plates in 5. Submedian plates on abdominal somites 3–5 bear- females better developed on left side and fused with ing concave, membranous areas. Abdominal somite marginal plates; marginal plates usually subdivided. 2 with a median plate and a marginal pair fused Sternite 5 (between pereopods 2) with deep median with the submedian plates. Rostrum more or less fissure. Scaphocerite absent or reduced to small spine. elongated, with two rounded terminal knobs...... Walking legs (pereopods 2–4) similar in form, third Phyllolithodes walking leg longest, always longer than pcl; dactyli of – Submedian plates on abdominal somites 3–5 lacking adults with or without corneous spines along flexor concave, membranous areas...... 6 margin.

6. Abdominal somite 2 composed of five distinct Composition plates. Marginal plates on abdominal somites 3–5 fused with the submedian plates ...... Rhinolithodes L. aequispina Benedict, 1895 [northern Pacific Ocean] L. aotearoa sp. nov. [New Zealand, southwestern Pacific – Abdominal somite 2 entire, formed by a single Ocean] plate...... 7 L. australiensis sp. nov. [Australia, southwestern Pacific 7. Rostrum thick, non-spiniform, hammer-shaped. Ocean] Scaphocerite small, rudimentary. Median plate on L. ceramensis Takeda & Nagai, 2004 [Indonesia, western abdominal somites 3–5 subdivided Sculptolithodes Pacific Ocean] – Rostrum more or less spiniform. Scaphocerite well- L. chaddertoni sp. nov. [Western Australia, eastern developed. Median plates on abdominal somites Indian Ocean] 3–5 entire...... 8 L. confundens Macpherson, 1988b [southwestern At- lantic Ocean] 8. Rostrum formed by a median spine and one dorsal L. couesi Benedict, 1895 [northern Pacific Ocean] spine or granule...... Glyptolithodes L. ferox Filhol, 1885b (= L. pyriformis Birstein & Vinogra- – Rostrum formed by a median spine and at least one dov, 1972) [Eastern Atlantic and Brazil] pair of dorsal spines...... 9 L. formosae Ahyong & Chan, 2010 [Taiwan, western Pacific Ocean] 9. Median plate on abdominal somite 3 subdivided L. galapagensis Hall & Thatje, 2009b [Galapagos Islands, into three. Walking leg 3 shorter than carapace eastern Pacific Ocean] width...... Lopholithodes L. jessica sp. nov. [New Zealand, southwestern Pacific – Median plate on abdominal somite 3 undivided. Ocean] Walking leg 3 equal to or longer than carapace L. longispina Sakai, 1971 [Japan, Western Pacific width...... Paralomis Ocean] L. macquariae sp. nov. [Macquarie Ridge, Southern Ocean] Lithodes Latreille, 1806 L. maja Linnaeus, 1758 [North Atlantic Ocean] L. mamillifer Macpherson, 1988c [southwestern Indian Lithodes Latreille, 1806: 39 [type species: Cancer maja Linnaeus, Ocean] 1758: 269. Gender: masculine]. L. manningi Macpherson, 1988b [western Atlantic Pseudolithodes Birstein & Vinogradov, 1972: 356 [type species: Ocean] Pseudolithodes zenkevitchi Birstein & Vinogradov, 1972 = Lithodes santolla (Molina, 1782). Gender: masculine]. L. megacantha Macpherson, 1991 [French Polynesia, central Pacific Ocean] Diagnosis. Carapace pyriform, not covering bases L. murrayi Henderson, 1888 [southwestern Indian of walking legs; regions indicated; gastric region el- Ocean] evated above other regions; cardiac region triangular, L. nintokuae Sakai, 1978 [northern-central Pacific separated from gastric region by deep groove; cervi- Ocean] cal groove shallow, indistinct. Rostrum comprised of L. panamensis Faxon, 1893 [eastern Pacific Ocean] median spine (usually distally bifid) and one or two L. paulayi Macpherson & Chan, 2008 [Guam, western pairs of dorsal spines; prominent ventral rostral spine Pacific]

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ L. rachelae sp. nov. [Great Australian Bight, Southern 3. Rostrum as long as pcl in specimens exceeding 75 Ocean] mm pcl [Vanuatu to southeastern Australia, pos- L. richeri Macpherson, 1990 [southwestern Pacific sibly Indonesia]...... L. richeri Macpherson, 1991 Ocean] – Rostrum not exceeding 0.7 pcl in specimens exceed- L. robertsoni sp. nov. [New Zealand, southwestern ing 75 mm pcl...... 4 Pacific Ocean] L. santolla (Molina, 1782) (= Pseudolithodes zenkevitchi 4. Dorsal carapace spines short, stout, about 0.1 pcl or Birstein & Vinogradov, 1972) [southwestern Atlan- less in specimens exceeding 50 mm pcl [Macquarie tic Ocean, southeastern Pacific Ocean] Ridge to Auckland Islands, New Zealand]...... L. turkayi Macpherson, 1988b [southwestern Atlantic ...... L. macquariae sp. nov. Ocean, southeastern Pacific Ocean] – Dorsal carapace spines long, slender, exceeding L. turritus Ortmann, 1892 [western Pacific Ocean] 0.3 pcl in specimens exceeding 50 mm pcl (though L. unicornis Macpherson, 1984 [Valdivia Bank, south- often about 0.1 pcl or less in specimens exceeding eastern Atlantic Ocean] 120 mm pcl)...... 5 L. wiracocha Haig, 1974 [eastern Pacific Ocean]. 5. Abdominal somite 2 marginal plates fused with Remarks. Prior to the present study, Lithodes included ‘central’ plate. Posterior branchial margin with 8–12 22 species. Macpherson (1988a) listed 15 species of spines or teeth [New Zealand]...... Lithodes worldwide of which seven occur in the At- ...... L. aotearoa sp. nov. lantic Ocean and eight in the Indo-Pacific. Since then, – Abdominal somite 2 marginal plates not fused with seven species of Lithodes have been described, one submedian plate, separated by suture. Posterior from the Indian Ocean and six from the Pacific Ocean: branchial margin with 5–8 spines or teeth...... 6 L. mamillifer Macpherson, 1988b [southwestern Indian Ocean], L. richeri Macpherson, 1990 [New Caledonia], 6. Male pereopod 4 merus length 1.3 pcl; merus length: L. megacantha Macpherson, 1991 [French Polynesia], height ratio exceeding 9 [southwestern Australia] L. ceramensis Takeda & Nagai, 2004 [Indonesia], ...... L. rachelae sp. nov. L. paulayi Macpherson & Chan, 2008 [Guam], L. gala- – Male pereopod 4 merus length less than 1.2 pcl; pagensis Hall & Thatje, 2009b [Galapagos Islands], and merus length:height ratio less than 8...... 7 L. formosae Ahyong & Chan, 2010 [off Dasi, northeastern Taiwan]. Four species are known from New Zealand to 7. Male pereopod 4 merus length up to 1.1 pcl in speci- Macquarie Island, all new, and four from around the mens exceeding 100 mm pcl. Posterior branchial Australian mainland, of which three are new species. margin with 7 or 8 spines or teeth [southeastern The results of the present study increase the number of New Zealand]...... L. robertsoni sp. nov. known species of Lithodes to 29. The diversity of Lithodes – Male pereopod 4 merus length less than 1.1 pcl in in the southwestern Pacific appears to be remarkably specimens exceeding 100 mm pcl. Posterior branchi- high compared to other regions, with six species from al margin with 5 or 6 spines or teeth [southeastern New Zealand and southeastern Australia alone. Seven Australia]...... L. australiensis sp. nov. species are known from the entire Atlantic Ocean, and a further 13 species are known from throughout the remainder of the Pacific Ocean. At present, only four species of Lithodes are known from the Indian Ocean, Lithodes aotearoa sp. nov. including L. chaddertoni sp. nov. and L. rachelae sp. nov. (Figs 5–12, Pl. 1A–B, 4C) described below. Lithodes murrayi. — Dell, 1963: 62. — Yaldwyn & Dawson, Key to adult Lithodes from New Zealand and 1970: 279–282, figs 1–3. — Dawson & Yaldwyn, 1985: 70 [New Zealand occurrences]. — McLay, 1988: 22. — Take- Australia da in Amaoka et al., 1990: 360. — Webber, 1997: 81, fig. 1. Flexor margin of walking leg dactyli lined with 4. — O’Shea et al., 1999: 49, fig. 15. — Zaklan, 2002: 766 small corneous spines...... 2 [New Zealand occurrences]. — Webber & Naylor, 2004a: 78–79. — Naylor et al., 2005: 41–42. — Ahyong et al., 2007: – Flexor margin of walking leg dactyli smooth...... 3 154. [Not L. murrayi Henderson, 1888]. 2. Chelae in both sexes equal, subcylindrical, non- Lithodes longispina. — Dawson & Yaldwyn, 1985: 70. — McLay, L longispina spinose [New Zealand]...... L. jessica sp. nov. 1988: 32 [Not . Sakai, 1971]. Lithodes spp. — O’Driscoll et al., 2003: 62. – Chelae dimorphic, palm highest distally, surfaces Lithodes cf. longispinus. — Webber & Naylor, 2004a: 79. spinose [Western Australia]...... — Naylor et al., 2005: 43–44. — Ahyong et al., 2007: 153...... L. chaddertoni sp. nov.

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 5. Lithodes aotearoa sp. nov., male holotype, cl 152.3 mm, pcl 88.6 mm, cw 85.3 mm, off Hawkes Bay (NIWA 34924). A, dorsal habitus. B, carapace, right lateral view.

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Type material. Holotype: NIWA 34924, male (cl 152.3 coll. M. Clark, 10 Dec 1985; NMNZ, 1 male (cl 69.0 mm, mm, pcl 88.6 mm, cw 85.3 mm), off Hawkes Bay, pcl 35.9 mm, cw 33.3 mm), 37°40.4–38.9’S, 179°25.7– 39°29.44–28.51’S, 178°25.05–25.48’E, 1000–980 m, 27.5’E, 911–1106 m, trawl, Trip 774, tow 15, coll. A. KAH9907/37, Z10169, 3 Jun 1999. France, 14 Jul 1994; NMNZ Cr11721, 1 male (pcl 118.7 Paratypes: NIWA 34907, 1 female (cl 96.0 mm, pcl mm, cw 114.6 mm), SE of East Cape, 37°53.9–57.6’S, 55.3 mm, cw 52.9 mm), off Hawkes Bay, 39°29.44– 178°55.8–56.8’E, 952–1052 m, FV Twofold Bay, coll. N. 28.51’S, 178°25.05–25.48’E, 1000–980 m, KAH9907/37, Bagley, 6 Jun 1985. Z10169, 3 Jun 1999; NMNZ Cr11738–11739, 1 ovigerous Hawkes Bay: NMNZ Cr11740, 1 female (pcl 131.4 female (pcl 125.1 mm, cw 125.7 mm), 1 female (cl 175.5 mm, cw 127.0 mm), Ritchie Bank, 39°29.8’S, 178°23.4’E, mm, pcl 130.8 mm, cw 127.7 mm), Hawke Seavalley, 900–947 m, orange roughy trawl, FV Arrow, A01/70/87, 39°38’S, 177°55’E, 1050 m, mud bottom, coll. G. Gibbs, 7 Jul 1987. 21 Sep 2001; NMNZ Cr11691, 1 male (cl 198.4, pcl 135.5 Louisville Ridge: NMNZ Cr11729, 1 ovigerous fe- mm, cw 133.3 mm), vicinity of Hawkes Bay, coll. A. male (cl 145.0 mm, pcl 96.2 mm, cw 84.8 mm), 40°58’S, Claudatus of Star Fish Supply Ltd, early Feb 1990; 165°04’W, 800 m, FV Peterson, coll. M. Miranovich, 2 NMNZ Cr11692, 1 male (cl 176.7, pcl 106.7 mm, cw Mar 1995. 102.7 mm), vicinity of Hawkes Bay, coll. A. Claudatus Wairarapa coast: NMNZ Cr14895, 1 juvenile female of Star Fish Supply Ltd, early Feb 1990. (cl 63.5+ mm, pcl 38.8 mm, cw 34.0 mm), 30.5 nautical miles [56 km] off Cape Turnagain, 1006–1189 m, FV Other material examined. West Norfolk Ridge: NMNZ Twofold Bay, A.D. Carruthers, 28 Dec 1984; NMNZ Cr11149, 1 male (cl 197.2 mm, pcl 145.7 mm, cw 143.3 Cr11727, 1 female (cl 170.0 mm, pcl 125.5 mm, cw mm), 34°19.14–17.19’S, 168°24.63–21.85’E, 842–831 m, 121.2 mm), off Cape Palliser, 40–41°30’S, 176°00–30’E, TAN0308/140, 2 Jun 2003. 803–1170 m, J18/-/84; NMNZ Cr12030, 1 male (pcl Challenger Plateau: NMNZ Cr11722, 1 male (cl 73.3 mm, cw 71.5 mm), off Wairarapa, 41°24.7–23.8’S, 190.5 mm, pcl 136.1 mm, cw 137.9 mm), 37°13.7’S, 176°08.4–12.1’E, 1103–1071 m, bottom trawl, RV James 167°49.9–51.9’E, 1109–1123 m, FV Arrow, A02/139/82, Cook, J18/4/84, 16 Oct 1984; NIWA 34920, 1 male (cl 9 Sep 1983; NMNZ Cr4699, 1 male (pcl 128.1 mm, 42.9, pcl 21.0 mm, cw 19.6 mm), 41°46.95–46.83’S, cw 127.0 mm), N Challenger Plateau, 37°19.7–20.0’S, 175°23.98–24.24’E, 1040–1053 m, TAN0616/79, 13 Nov 168°32.7–43.9’E, 1078–1082 m, orange roughy trawl, 2006; NIWA 29294, 1 damaged female (pcl 21.3 mm), bottom temperature 5.4°C, FV Albert Sandford, coll. 41°46.96–46.87’S, 175°23.98–24.13’E, 1053–1050 m, R. Stewart, 1 Oct 1986; NMNZ Cr11736, 1 female (cl TAN0616/83, 13 Nov 2006; NMNZ Cr11601, 1 male (cl 189.4, pcl 143.6 mm, cw 137.7 mm), W of Westport, 219.0 mm, pcl 163.7 mm, cw 165.9 mm), off Wairarapa 37°24.3’S, 168°59.0’E, 1020 m, FV Arrow, A4/137/83, coast, 700 m, D. Wylie, 17 Oct 1986; NMNZ Cr11584, 17 Oct 1983; NMNZ Cr11723, 1 male (crushed, pcl 1 ovigerous female (pcl 165.0 mm, cw 155.5 mm), off ~123 mm; with Briarosaccus callosus Boschma, Rhizo- Wairarapa coast, 700 m, coll. D. Wylie, 17 Oct 1986. cephala), 41°41.2–39.5’S, 168°44.2–44.8’E, 1043–1056 m, Marlborough coast: NMNZ Cr14896, 1 female (pcl FV Arrow, A03/48/83, 16 Oct 1983; NMNZ Cr4698, 1 19.2 mm, cw 17.3 mm), 1 crushed male (pcl approx. ovigerous female (pcl 161.3 mm, cw 156.0 mm), Chal- 18.0 mm), 35 km E of Clarence River mouth, 42°16.30’S, lenger Plateau, coll. R. Stewart, 30 Sep 1986. 174°20.80’E, 860–790 m, BS665, NZOI R23, “pink spines, Bay of Plenty: NMNZ Cr12036, 1 female (cl 104+ mm, legs and chelae, white body, black eyes”, 14 Jan 1979; pcl 69.6 mm, cw 68.5 mm), 37°22.80’S, 176°54.54’E, 1000 NIWA 34921, 1 ovigerous female (cl 164.3 mm, pcl 116.5 m, coll. G. Gibbs, 20 Nov 2001; NMNZ Cr14899, 1 male mm, cw 114.9 mm), 42°29.10–28.87’S, 173°36.91–36.28’E, (cl 41.7 mm, pcl 20.1 mm, cw 18.3 mm), N of Lottin 1079–1023 m, TAN0616/99, 15 Nov 2006. Point, 37°28.39–28.09’S, 178°21.00–19.80’E, 926–853 m, Chatham Rise: NMNZ Cr12029, 1 female (pcl 55.7 BS689 (NZOI R47), 18 Jan 1979; NIWA 42886, 1 male mm, cw 54.4 mm), NW of Chatham Islands, 42°42.0– (cl 194.8 mm, pcl 154.0 mm, cw 148.1 mm), 1 female (cl 41.7’S, 178°01.0–03.7’W, 1025–1055 m, COR/223/89, 13 189.8 mm, pcl 127.1 mm, cw 124.0 mm), Bay of Plenty, Aug 1989; NMNZ Cr11734, 1 male (pcl 66.9 mm, cw coll. Moana Pacific Fisheries, 2008; NMNZ, 1 female 64.9 mm), 42°44.3–44.0’S, 177°38.4–50.1’W, 1158–1121 (pcl 140.7 mm, cw 138.9 mm), Bay of Plenty, 37°17.5’S, m, Trip 2014, tow 131, J. Yeoman, 20 Feb 2005; NMNZ 176°58.67’E, 1000 m, crab pot, FV Savannah, coll. G. Cr6560, 1 male (pcl 88.7 mm, cw 86.5 mm), NW of Gibbs, 20 Nov 2001. Chathams, 42°46.2–46.5’S, 177°41.7–44.6’W, 988–993 East Cape: NMNZ Cr11714, 1 male (cl 182.7 mm, m, FV Cordella, COR/221/89, 13 Aug 1989; NIWA pcl 141.6 mm, cw 148.0 mm), off East Cape, 37°24.9’S, 34901, 1 male (cl 103.0 mm, pcl 58.1 mm, cw 57.7 179°10.0’E, 1014–1109 m, RV Wanaka, WK01/78/85, 5 mm), 42°47.17–45.66’S, 179°52.51–52.57’W, 978–1030 Jul 1985; NMNZ Cr4875, 1 male (cl 176.4 mm, pcl 131.5 m, TAN9908/25, Z10189, 21 Jun 1999; NIWA 34922, 1 mm, cw 129.7 mm), E of East Cape, 37°40.4–39.1’S, ovigerous female (cl 136.7 mm, pcl 99.0 mm, cw 96.3 179°11.4–10.1’E, 1123–1125 m, FV Wanaka, WK3/23/85, mm), 42°47.22–47.16’E, 176°43.33–42.66’W, 996–1009

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 6. Lithodes aotearoa sp. nov., male holotype, cl 152.3 mm, pcl 88.6 mm, cw 85.3 mm, off Hawkes Bay (NIWA 34924). A, carapace. B, left pereopod 4, dorsal. C, left pereopod 4 dactylus.

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ m, TAN0705/163, 16 Apr 2007; NIWA 33752, 1 male Chatham Islands, 44°33.65’S, 177°34.07’W, 895–910 (cl 212.0 mm, pcl 181.3 mm, cw 185.0 mm), 42°52.77– m, COR9004/152, FV Cordella, 21 Nov 1990; NIWA 52.93’S, 176°27.54–25.52’E, 838–844 m, TAN0709/6, 7 60555, 1 juvenile female (pcl 29.5 mm, cw 27.0 mm), Jul 2007; NIWA 42895, 1 male (pcl 148.0 mm, cw 149.9 44°39.75–40.42’S, 177°29.43–29.48’W, 1024–1043 m, FV mm), 42°58.89–58.15’S, 174°14.86–18.84’E, 988–993 m, San Waitaki, SWA0901/013, 6 Nov 2009; NIWA 34915, TAN0701/124, 20 Jan 2007; NIWA 48960, 1 male (pcl 1 female (pcl 87.8 mm, cw 85.8 mm), 44°48.75–48.49‘S, 190.4 mm, cw 196.1 mm), 43°31.1’S, 174°38.8’E, 512–547 172°32.2–35.02’E, 921–934 m, TAN9511/001, 5 Oct 1995; m, MFish Observer, trip 2814, tow 81, FV San Enter- NMNZ Cr9213, 2 juvenile females (pcl 29.2–32.1 mm, prise, coll. Ted Turton, 2009; NIWA 48961, 1 male (pcl cw 26.6–28.9 mm), 44°50.6’S, 175°29.4’E, 1044–1050 189.0 mm, cw 189.7 mm), 43°31.1’S, 174°38.8’E, MFish m, RV James Cook, JC090015/5, 20 Sep 1990; NMNZ Observer, trip 2814, tow 81, FV San Enterprise, coll. Ted Cr11145, 1 juvenile female (cl 54.5 mm, pcl 25.7 mm, cw Turton, 2009; NMNZ, 1 male (cl 198.2 mm, pcl 148.1 23.1 mm), 44°50.6’S, 175°29.4’E, 1044–1050 m, RV James mm, cw 152.6 mm), 44°00.5–00.8’S, 178°40.5–43.7’E, Cook, JC090015/5, 20 Sep 1990; NIWA 60530, 1 male (pcl 775 m, FV Kalinovo K01/063/81, 3 Dec 1981; NIWA 50.3, cw 49.3 mm), 42°48.89–48.33’S, 179°30.57–28.53’E, 48962, 1 male (cl 215.0 mm, pcl 178.9 mm, cw 184.5 1042–1049 m, TAN0509/11, 21 Jun 2005.. mm), 44°02’S, 174°46’E, MFish Observer, trip 2814, tow Stewart-Snares Shelf: NMNZ Cr1112, 1 male (cl 209.0 82, FV San Enterprise, coll. Ted Turton, 2009; NIWA mm, pcl 167.8 mm, cw 171.9 mm), SE of Solander Is- 61189, 1 ovigerous female (pcl 117.4 mm, cw 115.3 land, Foveaux Strait, in crayfish pot, 382 fathoms [699 mm), 44°03.78–03.39’S, 178°57.10–55.32’E, 757–765 m, m], coll. C. de Rich, Nov. 1960; NIWA 60557–60561, TAN9901, Z9615, 13 Jan 1999; NIWA 34612, 1 male (cl 2 males (pcl 123.4–143.0 mm, cw 124.8–141.9 mm), 3 128.4 mm, pcl 80.9mm, cw 79.2 mm), 44°05.73–06.63’S, ovigerous females (pcl 121.1–138.4 mm, cw 115.6–137.9 174°43.55–45.23’W, 1051–1020, TAN0709/109, 21 Jul mm), W of Stewart Island, 46°37.16’S, 166°37.80’E, 2007; NIWA 34898, 1 male (cl 81.4 mm, pcl 45.5 mm, cw 819–832 m, TAN0911/78, 15 Dec 2009. 43.6 mm), 44°06.49–07.69’S, 178°32.50–37.99’E, 906–926 Bounty Trough: NMNZ Cr11582, 1 male (cl 209.8 m, Z10853, 7 Apr 1999; NIWA 34905, 1 juvenile female mm, pcl 161.8 mm, cw 156.5 mm), Bounty Plateau, (pcl 31.9 mm, cw 29.4 mm), 44°06.58–06.81’S, 178°27.10– 47°47.02–49.65’S, 179°25.17–27.40’W, 596–562 m, 27.81’E, 963–955 m, TAN0011/043, 3 Nov 2000; NIWA TAN9310/001, 15 Nov 1993. 42894, 1 male (pcl 140.5 mm, cw 131.0 mm), 44°07.29– Bounty Plateau: NIWA 61188, 1 male (cl 226.3 mm, 07.99’S, 179°02.27–06.35’W, 361–364 m, TAN0701/62, pcl 183.7 mm, cw 180.0 mm), SE of Bounty Islands, 8 Jan 2007; NMNZ Cr11761, 1 female (cl 199 mm, 48°05.99’S, 179°56.99’W, 252 m, FIS Endeavour, F122, pcl 168.0 mm, cw 161.0 mm), 44°0.15’S, 178°31.25’E, 26 Jan 1965; NMNZ Cr11583, 1 male (cl 194.0 mm, pcl 775–790 m, FV Kalinovo, K01/064/81; NMNZ Cr6561, 157.4 mm, cw 162.2 mm), 48°20.74–21.58’S, 179°53.71– 1 male (pcl 138.8 mm, cw 140.4 mm), W of Chathams, 57.42’W, 538–541 m, RV Tangaroa, TAN9310/002, 15 44°17.2–17.4’S, 179°56.0–59.2’W, 828–840 m, FV Cord- Nov 1993; NMNZ Cr11733, 1 male (cl 165.9 mm, pcl ella, COR/204/89, 9 Aug 1989; NIWA 60582, 1 male (pcl 117.0 mm, cw 113.4 mm), Bounty Plateau, 48°22.5’S, 179.2 mm, cw 178.5 mm), 44°22.83–22.66’S, 179°54.68– 179°56.0’E, 547–561 m, RV Shinkai Maru, Cruise II, 56.27’E, 952–957 m, FV San Waitaki, SWA0901/82, 16 prob. Stn 2/80, 17 Nov 1975; NIWA 60616, 1 juvenile Nov 2009; NIWA 61159, 1 juvenile male (pcl 42.2 mm, female (cl 32.3 mm, pcl 13.4 mm, cw 10.8 mm), NE of cw 41.1 mm), 44°22.83–22.66’S, 179°54.68–56.27’E, Antipodes Islands, 49°18.63–18.06’S, 179°47.88–48.33’E, 952–957 m, FV San Waitaki, SWA0901/82, 16 Nov 2009; 1506–1476 m, TAN0307/59, 29 Apr 2003. NIWA 60566, 1 male (cl 206.2 mm, pcl 169.0 mm, cw Campbell Plateau: NIWA 42799, 1 male (pcl 102.5 171.1 mm), 44°25.78’S, 178°42.07–41.67’W, 697–812 m, mm, cw 97.4 mm), 46°57.58–57.04’S, 170°11.11–15.43’E, FV San Waitaki, SWA0901/74, 15 Nov 2009; NIWA 855–946 m, TAN9805/02, 7 Apr 1998; NIWA 60565, 1 60567, 1 female (cl 230.5 mm, pcl 190.5 mm, cw 183.9.1 male (pcl 151.0 mm, cw 150.5 mm), 1 female (pcl 128.6 mm), 44°25.78’S, 178°42.07–41.67’W, 697–812 m, FV San mm, cw 121.8 mm; with Briarosaccus callosus Boschma, Waitaki, SWA0901/74, 15 Nov 2009; NIWA 42893, 1 Rhizocephala), 48°13.47’S, 169°53.74’E, 802–831 m, ovigerous female (cl 158.0 mm, pcl 118.9 mm, cw 116.0 TAN0911/5, 28 Nov 2009; NIWA 60551, 1 male (cl mm), 44°26.80–26.73’S, 175°37.05–32.85’E, 760–779 m, 240.0 mm, pcl 195.0 mm, cw 199.1 mm), Pukaki Rise, TAN0701/86, 13 Jan 2007; NMNZ Cr11693, 1 male 48°45.42’S, 172°35.90’E, 700–717 m, TAN0911/11, (pcl 170.0 mm, cw 173.3 mm), 44°27.9’S, 173°40.3’E, 20 Nov 2009; NIWA 34891, 1 male (cl 233.8 mm, pcl 727 m, FV Shinkai Maru, net 32, 8 Mar 1983; NIWA 194.0 mm, cw 198.6 mm), 49°15.99’S, 168°42.99’E, 785 60553, 1 ovigerous female (cl 191.4 mm, pcl 145.4 mm, m, Z9312, 15 Sep 1998; NMNZ Cr4858, 1 female (cl cw 137.0 mm), 1 female (cl 141.1 mm, pcl 99.2 mm, cw 81+ mm, pcl 64.4 mm, cw 61.9 mm), SW of Urry Bank, 98.8 mm), 44°32.78–33.62’S, 175°23.73–23.77’S, 939–985 49°30.6–39.54’S, 174°08.6–01.8’E, 798 m, FV Oyang, #7, m, SWA0901/61, FV San Waitaki, 13 Nov 2009; NMNZ Tow 5, coll. C.D. Roberts, 7 May 1987; NMNZ Cr11694, Cr14900, 1 female (pcl 28.7 mm, cw 26.3 mm), SW of 1 male (pcl 106.6 mm, cw 108.6 mm), SE of Campbell

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 7. Lithodes aotearoa sp. nov., male holotype, cl 152.3 mm, pcl 88.6 mm, cw 85.3 mm, off Hawkes Bay (NIWA 34924). A, right chela. B, left chela. C, carapace, posterior surface. D, right antenna. E, abdomen. F, abdominal somite 2.

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 8. Lithodes aotearoa sp. nov., ovigerous female, pcl 125.5 mm, off Cape Palliser (Cr11727). A, dorsal habitus. B, abdomen. 22

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 9. Lithodes ?aotearoa sp. nov., aberrant ovigerous female, cl 145.0 mm, pcl 96.2 mm, cw 84.8 mm, Louisville Ridge (Cr11729). A, dorsal habitus. B, carapace. C, abdomen. 23

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Island, 53°S, 170°E, 690–875 m, stn 139 & 140, FV region with pair of anterior tubercles or small spines Wessermunde, 13 May 1979; NMNZ Cr11695–11696, 1 and posterior pair of long upright spines. Hepatic male (pcl 87.7 mm, cw 83.6 mm), 1 female (pcl 80.5 mm, spine longest of carapace spines (0.6 pcl in holotype), cw 81.1 mm), SE of Campbell Island, 53°46.7–48.5’S, directed anterolaterally. Intestinal region with pair of 168°27.5–35.0’E, 870–880 m, trawl, FV Wessermunde, stn long upright spines on posterior margin. Branchial 136, 12 May 1979; NMNZ Cr7352, 1 male (cl 206.8 mm, surface with 2 or 3 long upright spines as follows: 1 pcl 153.3 mm, cw 156.5 mm), SE of Campbell Island, spine at level of pereopod 3 coxa, 1 or 2 in transverse 53°28.0’S, 171°22.2’E, 420 m, FV Ryuyo Maru, JQET, tow row at level of pereopod 4 coxa, innermost varying 168, coll. H. Kavale, 30 Aug 1987. from a spiniform tubercle to as long as outer spine, New Zealand, no specific locality: NMNZ Cr11728, 1 outer spine often with short spine at base. Branchial male (pcl 170.0 mm, cw 178.0 mm), between Ninety margins spinose; anterior branchial margin with 2 Mile Beach and Wairarapa, RV Kalinovo, K01/?/81, spines, posterior spine longer than anterior; 1st primary 1981; NMNZ Cr14898, 1 male (pcl 31.7 mm, cw 29.4 marginal branchial spine subequal to or shorter than mm), SE New Zealand, from MAF, 1985; NIWA 34904, hepatic spine; lateral branchial margin with 2 or 3 short 1 female (cl 82.3 mm, pcl 40.8 mm, cw 37.2 mm), New spines; 2nd primary marginal branchial spine quarter to Zealand, MAF, 16 Jan 1989; NMNZ Cr11252, 1 male (cl half length of hepatic spine; posterior branchial margin 178.7 mm, pcl 140.0 mm, cw 129.4 mm; with Briarosac- with 8–12 spines. cus callosus Boschma, Rhizocephala), New Zealand, Pterygostomian region granulate or tuberculate, no other data; NMNZ Cr11253, 1 female (pcl 154.3 with small, anterior, submarginal spine. mm, cw 132.8 mm; with Briarosaccus callosus Boschma, Rostrum 0.18–0.71 pcl (becoming proportionally Rhizocephala), New Zealand, no other data; NMNZ shorter with increasing size), comprising proximal and Cr11735, 1 male (cl 185.6 mm, pcl 143.0 mm, cw 146.3 distal portions; proximal portion angled dorsally by at mm), floating off Wellington wharf, coll. J. Annala, least 30°, with pair of long, divergent, dorsal spines that Dec 1985. overreach midlength of distal portion of rostrum; distal portion subhorizontal, distally bifurcate for one-third Diagnosis. Carapace dorsal surface with slender spines to two-thirds length, forming pair of strongly divergent becoming reduced to short conical spines above about spines; ventral rostral spine elongate, extending ante- 120 mm pcl (usually about 0.1 pcl); gastric region with riorly beyond apices of outer orbital spine. Posterior 4 upright spines, anterior pair longer than half length orbital margin concave; outer orbital spine directed of hepatic spine, posterior pair half length of hepatic anteriorly, about as long as anterolateral spine, reach- spine; cardiac and intestinal regions each with 2 long ing beyond cornea. upright spines; branchial surface with 2 or 3 long up- Ocular peduncle: Longer than cornea; unarmed or right spines. Branchial margins spinose; 1st primary with indistinct, scattered granules. marginal branchial spine subequal to or shorter than Antennule: Peduncle unarmed, reaching anteriorly hepatic spine, 2nd primary marginal branchial spine beyond antennal peduncle by about half length of distal about half length of hepatic spine or less; posterior antennular peduncle article. branchial margin with 8–12 spines. Rostrum not ex- Antenna: basal antennal article unarmed; outer mar- ceeding 0.7 pcl; proximal half strongly upraised by gin of article 2 with long slender spine, not reaching 30–45°. Abdominal somite 2 comprising single plate beyond proximal quarter of article 5; article 3 unarmed; in adults. Antennal peduncle article 2 with outer spine scaphocerite minute, blunt, shorter than article 4 (or not reaching midlength of article 4. Chelipeds unequal; produced to a slender spine on right side in one speci- palms with low tubercles or spines on dorsal surface. men, male, NMNZ Cr11735); article 4 unarmed, about Walking legs spinose; surface between major spines half as long as article 5. smooth or with very few, scattered spines. Pereopod Abdomen: Somite 1 with triangular teeth on posterior 4 merus about as long as or shorter than pcl, length margin. Somite 2 sparsely granulate, submedian plates about 7 times height or less. Dactylus 0.5–0.7 propodus fused with median by 40–60 mm pcl; median surface length; flexor margin unarmed. with pair of slender submedian spines; posterior margin with 6 stout spines or angular prominences; lateral Description. Carapace: Pyriform, 0.97–1.14 times longer margins irregularly dentate. Somites 3–5 with nodular than wide; regions indicated; dorsal surface armed with median plates; surface of submedian plates irregular, long, slender spines (generally reduced to short conical nodular, with scattered acute tubercles; marginal plates spines of about 0.1 pcl above about 120 mm pcl) and of males subdivided, with angular or sharp apices; scattered, widely separated granules or minute second- marginal plates of females with left margin crenulate ary spines, surface otherwise smooth. Gastric region to spinose (fused with submedians), right marginals convex, with 2 pairs of long, upright spines. Cardiac subdivided, with angular to spiniform apices. Somite

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 10. Lithodes aotearoa sp. nov. A–D, juvenile female, pcl 25.7 mm, Chatham Rise (NMNZ Cr11145). E–F, female, pcl 69.6 mm, Bay of Plenty (NMNZ Cr12036). A, E, dorsal view. B, F, lateral view. C, ventral view. D, abdominal somite 2.

6 of both sexes as long as or longer than wide, with 2 covered with granules in addition to major spines. proximal and 4 distal spines or prominences. Telson Coxa with blunt tubercles and tufts of setae, unarmed; rounded, unarmed, at most with 4 blunt granules. ischiobasis with 3 stout ventral spines and about 6 blunt Pereopod 1 (chelipeds): Dimorphic but spination tubercles. Merus inner margin, with slender subdistal similar. Surfaces of both chelipeds smooth or sparsely spine and 2 or 3 smaller spines; ventral margin with

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ two rows of 2 or 3 low conical spines; dorsal and lateral (male), 0.84–0.96 (female); length:height ratio 8.42– surface spinose, spines largest distally. Carpus with 12.29 (male), 7.17–10.62 (female). Dactylus 0.52–0.66 prominent spines on dorsal and lateral surfaces; dor- propodus length (male), 0.53–0.64 (female). sal margin with row of 4 stout spines; lateral margin Pereopod 3 length 2.81–2.66 pcl (male), 1.89–2.72 with 4 or 5 stout spines of similar size to dorsal row; pcl (female). Merus 0.94–1.05 pcl (male), 0.63–0.96 pcl ventral margin small scattered acute tubercles. Palms (female); length:height ratio 5.72–7.34 (male), 4.20–6.35 of both chelipeds in both sexes with similar ornamenta- (female). Carpus 0.50–0.53 merus length (male), tion; with small spines and acute tubercles on dorsal, 0.50–0.61 (female). Propodus 0.79–0.88 merus length lateral and ventral surfaces, inner surface with acute (male), 0.86–0.92 (female); length:height ratio 8.74– tubercles; dorsal margin with row of about 5 small 11.45 (male), 7.85–10.77 (female). Dactylus 0.50–0.60 conical spines; midlateral surface with 2 rows of 3 or propodus length (male), 0.54–0.61 (female). 4 small spines, of similar size to dorsal row; ventral Pereopod 4 length 2.68–2.98 pcl (male), 1.89–2.77 surface with scattered tubercles, smaller than lateral pcl (female). Merus 0.91–1.06 pcl (male), 0.62–0.95 pcl and dorsal spines. (female); length:height ratio 5.56–7.16 (male), 4.55–6.43 Major cheliped 1.17–1.45 pcl (male), 1.09–1.36 pcl (female). Carpus 0.52–0.55 merus length (male), (female); upper palm length 0.96–1.39 times height 0.53–0.64 (female). Propodus 0.87–0.92 merus length (male), 1.14–1.48 (female); occlusal margins of fingers (male), 0.89–0.96 (female); length:height ratio 9.12– corneous for distal third, proximally with 3 calcareous 13.04 (male), 8.56–10.41 (female). Dactylus 0.49–0.66 nodules; dactylus dorsal margin broadly convex, with propodus length (male), 0.57–0.62 (female). rows of tufts of golden setae and blunt proximal tooth, Egg size: 2.2–2.3 mm diameter. 1.21–1.42 times longer than dorsal margin of palm (male), 1.25–1.62 (female). Colour in life. Usually deep-purplish red overall, whit- Minor cheliped 1.22–1.36 pcl (male), 0.96–1.35 pcl ish along cervical groove, often with whitish or pinkish (female); upper palm length 1.21–1.53 times height blotches on pereopods and carapace; occasionally pale (male), 1.19–1.57 (female); occlusal margin corneous pinkish-red or whitish overall in largest specimens for slightly less than distal half, proximally crenulate; (Pl. 1A, B). dactylus dorsal margin broadly convex, with rows of tufts of golden setae and blunt proximal tooth, 1.61– Etymology. Named aotearoa, for the occurrence of the 1.72 times longer than dorsal margin of palm (male), species in New Zealand; used as a noun in apposi- 1.59–1.83 (female). tion. Pereopods 2–4 (walking legs 1–3): Similar, slender, elongate; ornamentation similar in both sexes; seg- Remarks. For several decades, Lithodes murrayi Hend- ments spinose, surface between major spines smooth erson, 1888 (type locality: Prince Edward Island, or with scattered spines or granules. Pereopod 4 long- southwest Indian Ocean) and L. longispina Sakai, 1971 est. Distal margins of coxae with low, blunt tubercles. (type locality: Japan) were believed to occur in New Ischiobasis with 3 or 4 non-setose distal spines. Merus Zealand waters (Dell 1963; Yaldwyn & Dawson 1970; subcircular to ovate in cross section; extensor margin Dawson & Yaldwyn 1985). Examination of collections with 8–10 spines of which 2 or 3 are distinctly longer on which New Zealand records of both species were in addition to long, prominent, distal spine; flexor based, however, revealed that neither L. murrayi nor margin with 2 rows of 5–8 spines; upper surface with L. longispina occurs in the region. Moreover, the New row of 6–8 spines. Carpus extensor margin with distal Zealand specimens previously reported as L. murrayi and second proximal spines longest, exceeding two- and L. longispina are referable to a single undescribed thirds carpus length; surface scattered with 8 or 9 small species, L. aotearoa sp. nov. The initial New Zealand spines. Propodus ovate in cross section; with 7–9 spines record of L. murrayi was based on a large specimen (pcl on extensor margin and 6–8 spines on dorsal surface; 167.8 mm, NMNZ Cr1112) from Foveaux Strait with flexor margin with 6 or 7 smaller spines. Dactylus relatively short dorsal spines (Dell 1963; Yaldwyn & curved, rounded in cross section distally; proximally Dawson 1970), though still considerably longer than with 2 spines on either side adjacent to articulation; those of the much smaller type specimens of L. mur- extensor margin with 1 or 2 small spines ; flexor mar- rayi from the southwestern Indian Ocean (pcl 64–72 gin smooth, unarmed, with few widely spaced setae; mm). New Zealand records of L. longispina were based apex corneous. on specimens smaller than 120 mm pcl in which the Pereopod 2 length 2.34–2.66 pcl (male), 1.78–2.54 dorsal spines are greatly elongate as in similarly sized pcl (female). Merus 0.83–0.96 pcl (male), 0.58–0.90 pcl Japanese specimens of L. longispina. Over the past two (female); length:height ratio 5.21–7.22 (male), 3.77–6.21 decades, numerous New Zealand specimens of Lithodes (female). Carpus 0.50–0.55 merus length (male), became available through several sources including 0.51–0.62 (female). Propodus 0.78–0.87 merus length commercial fishermen, fishery surveys and other deep-

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 11. Lithodes aotearoa sp. nov. A, male, pcl 80.9 mm, Chatham Rise (NIWA 34612). B, male paratype, pcl 106.7 mm, off Hawkes Bay (NMNZ Cr11692). C–D, male, pcl 163.7 mm, off Wairarapa coast (NMNZ Cr11601). E–F, male, pcl 167.8 mm, SE of Solander Island, Foveaux Strait (NMNZ Cr1112). A–C, E, dorsal habitus. D, F, carapace, right lateral view.

water research programmes. The difficulty of assigning species of Lithodes and Neolithodes, the proportional ‘intermediate-sized’ New Zealand specimens to either length of the dorsal spines is inversely related to body L. murrayi or L. longispina has been acknowledged (e.g. size. Failure to account for allometry by previous Webber & Naylor 2004a), but the authority afforded to workers has prevented recognition that New Zealand the original identifications of New Zealand specimens records of L. longispina and L. murrayi were based on has been such that their veracity has not been reas- phases of L. aotearoa. Despite significant allometric sessed until now. As in other lithodids, particularly variation in lithodids, spine length at any given size

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ varies between species. In L. longispina and L. aotearoa, pcl (NIWA 34907), 0.6 pcl in the 88.6 mm pcl holotype, long dorsal spines persist into maturity, though in the and 0.1 pcl in the 145.7 mm specimen (NMNZ Cr11149). largest specimens, dorsal spines are correspondingly Apart from reduction in spine size, other allometric short. Thus, spine length remains a very useful diag- changes include fusion of the median and marginal nostic character providing body size is considered. plates of abdominal somite 2 (fused by 40–60 mm pcl), Lithodes murrayi, a ‘short-spined’ species, attains little the flexor spination of the dactyli of the walking legs more than half the size of L. aotearoa (pcl 108 mm versus (present in juveniles up to about 20 mm pcl), the rela- 194 mm), with spines always considerably shorter and tive length of the walking legs (proportionally longer stouter than those of the latter at any given size. The with increasing size), and stoutness of the meri of the carapace spines of L. aotearoa are of similar form to adult walking legs (becoming less slender with increasing L. murrayi only in specimens exceeding 170 mm pcl, size). In addition, leg length and meral stoutness is well above the maximum size of the latter. Apart from sexually dimorphic. At any given size, females have dorsal spine shape and length, L. murrayi is readily proportionally shorter walking legs and longer cor- distinguished from L. aotearoa in bearing 5 or 6 versus responding meri than males (by about 15–30%). 8–12 posterior branchial marginal spines. In the majority of specimens, including most adults, Lithodes aotearoa closely resembles L. australien- the rostral apex is deeply bifurcate. In several speci- sis sp. nov. and L. rachelae sp. nov. from Australia, mens, however, the apex is shallowly bifurcate as in L. robertsoni sp. nov. from New Zealand, and to a lesser L. murrayi, and in four specimens (NIWA 33752, 60557, extent L. longispina from Japan, with which the first two 60567; NMNZ Cr11761), the apex is unilobate as in the new species were previously misidentified (Dawson & holotype of L. unicornis Macpherson, 1984. Four speci- Yaldwyn 1985; McLay 1988). The five species share the mens (NMNZ Cr11723, Cr11252–11253, NIWA 60565) numerous, greatly elongated spines on the carapace were infected by the rhizocephalan parasite, Briarosac- and pereopods in specimens up to about 120 mm pcl, cus callosus. In all but one specimen, the scaphocerite and the strongly upraised proximal portion of the is obsolete, represented by a small, blunt nodule. In a rostrum. The new species differs from L. longispina in 142.8 mm pcl male (NMNZ Cr11735), the scaphocerite bearing more pronounced and more numerous cara- is produced to a sharp point on the left side, and de- pace spines, especially on the branchial surface and veloped as a slender spine on the right side. margins: these differences are best observed in speci- An ovigerous female specimen from the Louisville mens of 100 mm pcl or less. Lithodes aotearoa bears 8–12 Ridge (NMNZ Cr11729, Fig. 9) differs from other posterior branchial marginal spines versus versus 5 or specimens of L. aotearoa in having three separate plates 6 in L. longispina. The branchial surfaces of L. aotearoa on abdominal somite 2 (rather than being fused into a bear two or three long dorsal spines of similar length single plate), seven posterior branchial marginal spines, to the cardiac spines: one anterior spine at the level more prominently spinose margins of the abdomen of the second walking leg, and one or two posterior and proportionally longer dactyli of the walking legs spines at the level of the third walking leg. In contrast, (0.7 versus 0.5–0.6). In the three-segmented abdominal L. longispina bears only the long anterior branchial somite 2 and posterior branchial marginal spines, the spine, with the posterior branchial spines replaced by specimen resembles L. robertsoni sp. nov. Apart from short acute tubercles. In specimens up to 100 mm pcl, the long walking leg dactyli, however, limb morphom- both L. aotearoa and L. longispina bear a long hepatic etry of the Louisville Ridge specimen agrees well with spine, two long primary marginal branchial spines, L. aotearoa, and thus differs from L. robertsoni, and and a pair of long intestinal spines. In L. longispina, the L. australiensis. The aberrant Louisville Ridge speci- anterior branchial margin is unarmed or bears only men could represent an undescribed species but until one or two minute spines, and the posterior branchial further material becomes available, the specimen is margin is armed with not more than six stout, well- tentatively assigned to L. aotearoa. spaced spines or tubercles of which two or three are Lithodes aotearoa is the largest species of lithodid in longer than the remainder. In specimens of L. aotearoa New Zealand waters, attaining carapace dimensions exceeding 120 mm pcl, the major carapace spines are of at least cl 240.0 mm, pcl 195.0 mm and cw 199.1 mm, reduced in size, although the positions and numbers with a leg span of at least 1.3 m (NIWA 60551). Fe- of spines are consistent with that of smaller specimens. males are ovigerous by at least 116.5 mm pcl (NIWA Features distinguishing L. aotearoa from L. rachelae, 34921), but appear to be mature by about 90 mm pcl L. australiensis, and L. robertsoni are outlined under the based on abdomen development and the presence of respective accounts of those species. coxal setae. The dorsal spine length in L. aotearoa becomes proportionally shorter with increasing body size. The Distribution. New Zealand, from the southern West longest carapace spine (hepatic) is longer than the pcl Norfolk Ridge south to The Snares, including the at 17.3 mm pcl (NMNZ Cr14896), 0.9 pcl at 55.3 mm Challenger Plateau, Bay of Plenty, Chatham Rise and

Campbell Plateau; 252–1506 m, usually 950–1100 m. Type material. (All New South Wales, Australia). Holo- An underwater image of a Lithodes sp. taken at 1242 type: AM P35607, male (cl 161.9 mm, pcl 112.2 mm, cw m at Rumble II seamount on the southern Kermadec 107.9 mm), E of Brush Island, 35°30’S, 150°52’E, 1065 m, Ridge, appears to be L. aotearoa (see cover). FRV Kapala, K83-19-12, coll. K. Graham, 6 Dec 1983. Paratypes: AM P80452, 1 male (cl 76.5 mm, pcl 39.0 Lithodes australiensis sp. nov. mm, cw 34.8 mm), 2 females (pcl 18.8–31.6 mm, cw 15.8–28.3 mm), off Cape Hawke, 32°08’S, 153°09’E, (Figs 13–17, Pl. 1C) 1007–1053 m, FRV Kapala, K84-10-04, 18 Jul 1984; AM Lithodes longispina. — Dawson & Yaldwyn, 1985: 70. — Poore P35601, 1 male (cl 166.0, pcl 97.0 mm, cw 90.7 mm), E et al., 1998: 77. — Davie, 2002: 72–73 [Australian occur- of Newcastle, 32°57–54’S, 152°44–46’E, 988–1033 m, rences]. — Poore, 2004: 266, 268, pl. 16c. [Not L. longispina FRV Kapala, K83-13-04, coll. K. Graham, 19 Oct 1983; Sakai, 1971] AM P81160, 1 male (pcl 40.0 mm, cw 35.5 mm), E of Shoalhaven Heads, 34°51–55’S, 151°15–14’E, 1042–1061 m, FRV Kapala, K83-14-05, coll. K. Graham, 26 Oct 1983;

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 13 (above). Lithodes australiensis sp. nov., male holotype, cl 161.9 mm, pcl 112.2 mm, cw 107.9 mm, E of Brush Island (AM P35607). A, dorsal habitus. B, carapace, right lateral view. Figure 14 (opposite). Lithodes australiensis sp. nov., male holotype, cl 161.9 mm, pcl 112.2 mm, cw 107.9 mm, E of Brush Island (AM P35607). A, right chela. B, left chela. C, carapace. D, right orbit and antenna. E, abdominal somite 2. F, abdomen.

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ AM P35595, 1 male (pcl 50.5 mm, cw 45.5 mm), E of intestinal regions each with 2 long upright spines; Shoalhaven Heads, 34°53–56’S, 151°15–13’E, 1079–1116 branchial surface with 2 long upright spines. Branchial m, FRV Kapala, K83-14-06, coll. K. Graham, 26 Oct 1983; margins spinose; 1st primary marginal branchial spine AM P80453, 1 male (cl 50.7 mm, pcl 23.9 mm, cw 20.6 shorter than two-thirds length of hepatic spine, 2nd mm), 2 females (cl 53.1–53.2 mm, pcl 24.4–26.0 mm, primary marginal branchial spine shorter than 1st cw 21.5–21.6 mm), E of Shoalhaven Heads, 34°53–56’S, primary marginal branchial spine; posterior branchial 151°15–13’E, 1079–1116 m, FRV Kapala, K83-14-06, coll. margin with 5 or 6 short spines and 1 or 2 low tubercles. K. Graham, 26 Oct 1983; AM P35599, 1 male (cl 199.7 Rostrum not exceeding 0.8 pcl; proximal half strongly mm, pcl 132.8 mm, cw 127.5 mm), NE of Brush Island, upraised by 30–45°. Abdominal somite 2 comprising 3 35°29’S, 150°52’E, 942 m, FRV Kapala, K84-04-03, coll. K. plates in juveniles through adults. Antennal peduncle Graham, 3 Apr 1984; AM P35591, 1 male (pcl 39.1 mm, article 2 with outer spine not reaching beyond article cw 36.3 mm), E of Gabo Island, 37°43–41’S, 150°20’E, 4. Chelipeds unequal; palms with low tubercles or 961–979 m, FRV Kapala, K83-12-01, coll. K. Graham, spines on dorsal surface. Walking legs spinose; sur- 26 Sep 1983. face between major spines smooth or with very few, scattered spines. Pereopod 4 merus shorter than 1.2 Other material examined. Tasmania: TM G2970, 1 female pcl (1.1 to almost 1.2 in males, 0.9 to as long as pcl in (pcl 38.0 mm, cw 35.4 mm), off Bicheno, E Tasmania, females), length less than 8 times height; propodus 1000 m, coll. G. Darcey, 25 Apr 1982; TM G3162, 1 length 12–13 times height (male), 11 (female); dactylus male (cl 137.0 mm, pcl 77.6 mm, cw 70.2 mm), ENE flexor margin unarmed. of St Patricks Head, 41°34.7–34.4’S, 148°44.6–46.6’E, 1090–1150 m, RV Soela stn 16, 9 May 1987; TM G3479, Description. Carapace: Pyriform, 1.01–1.14 times longer 1 crushed exuvium, E Tasmania, 41°39’S, 148°39’E, than wide; regions indicated; dorsal surface armed 915–970 m, 27 Jul 1982; TM G3490, 1 female (cl 71.3 with long, slender spines and scattered, widely sepa- mm, pcl 35. 6 mm, cw 34.8 mm), 15 miles [24 km] N rated granules. Gastric region convex, with 2 pairs of St Helens Hill, eastern Tasmania, 41°00’S, 148°46’E, of long, upright spines. Cardiac region with pair of 540–720 m, FV Ocean Raider, coll. D. Whennan, 13 Mar anterior tubercles and posterior pair of long upright 1991; NMV J55658, 2 males (pcl 25.9–30.6 mm, cw spines. Hepatic spine longest of carapace spines (0.26 24.5–28.0 mm), Southern Ocean, S of Tasmania, Pedra pcl in holotype), directed anterolaterally. Intestinal site, 44°15.67–15.98’S, 147°05.83–05.52’E, 730–1000 m, region with pair of long upright spines on posterior coll. T. O’Hara & T. Costa, 2 Apr 2007; TM G3582, 1 margin. Branchial surface with 2 long upright spines male (cl 47.7 mm, pcl 24.1 mm, cw 22.1 mm), Main and 2 conical tubercles as follows: 1 spine at level of Pedra, approx. 77 km SSE of Southeast Cape, 44°16’S, pereopod 3 coxa, 1 spine at level of pereopod 4 coxa, 2 147°04’E, 1110 m, epibenthic sled, RV Southern Sur- conical tubercles (or short spines in juveniles) in trans- veyor, SS01/97/06, 21 Jan 1997; TM G3659, 1 male verse row slightly behind posterior spine. Branchial (pcl 60.3 mm, cw 54.4 mm), Main Pedra, approx. 77 margins spinose; anterior branchial margin with 2 km SSE of Southeast Cape, 44°16’S, 147°08’E, 1312 m, small spines, posterior spine longer than anterior; 1st crab trap, RV Southern Surveyor, SS01/97/08, 21–25 primary marginal branchial spine shorter than two- Jan 1997; NIWA 41384, 1 juvenile female (cl 95.6, pcl thirds length of hepatic spine; lateral branchial margin 53.8 mm, cw 49.3 mm), South Tasman Rise, 47°09.0– with 1 short spine and 1 or 2 granules; 2nd primary mar- 27.99’S, 148°44.6–52.99’E, 1050 m, FV Arrow, 1164/4, ginal branchial spine shorter than 1st primary marginal Z9395, coll. C. Rattray, 9 Oct 1998; NIWA 34906, 1 branchial spine (usually about half length); posterior male (pcl 40.0 mm, cw 36.1 mm), South Tasman Rise, branchial margin with 5 or 6 short spines and 1 or 2 47°27.09–27.99’S, 148°53.01–52.99’E, 1012–1058 m, low tubercles. Pterygostomian region granulate or 1137/12, Z9231, 13 Aug 1998; NIWA 34899, 1 male (cl tuberculate, with small, anterior, submarginal spine. 89.7 mm, pcl 52.9 mm, cw 46.2 mm), South Tasman Rostrum as long as pcl in smallest specimens to Rise, 47°28.02–27.99’S, 148°53.05–53.05’E, 1031–1034 m, half pcl in largest specimens, comprising proximal and Z9232, 7 Aug 1998; NIWA 34903, 1 damaged male (pcl distal portions; proximal portion angled dorsally by at 78.6 mm, cw 73.3 mm), South Tasman Rise, 49°22.6’S, least 30°, with pair of long, divergent, dorsal spines that 150°27.0’E, 910–1200 m, Z10303. overreach midlength of distal portion of rostrum; distal portion subhorizontal, distally bifurcate for about Diagnosis. Carapace dorsal surface with slender spines half length, forming pair of strongly divergent spines; becoming reduced to short conical spines above about ventral rostral spine elongate, extending anteriorly 120 mm pcl; gastric region with 4 upright spines, beyond apices of outer orbital spine. Posterior orbital anterior pair longer than half length of hepatic spine, margin concave; outer orbital spine directed anteriorly, posterior pair half length of hepatic spine; cardiac and

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 15. Lithodes australiensis sp. nov., female, pcl 53.8 mm, S of Tasmania (NIWA 41384). A, dorsal habitus. B, abdomen.

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 16. Lithodes australiensis sp. nov. A, male, pcl 24.1 mm, SSE of Southeast Cape, Tasmania (TM G3582). B, male, pcl 60.3 mm, SSE of Southeast Cape, Tasmania (TM G3659).

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ slightly longer than anterolateral spine, reaching to or (male), 1.69–1.77 (female); occlusal margin corneous beyond cornea. for slightly less than distal half, proximally crenulate; Ocular peduncle: Longer than cornea; unarmed. dactylus dorsal margin broadly convex, with rows of Antennule: Peduncle unarmed, reaching anteriorly tufts of golden setae and blunt proximal tooth, 1.51– beyond antennal peduncle by about three-quarters 1.69 times longer than dorsal margin of palm (male), length of distal antennular peduncle article. 1.51–1.56 (female). Antenna: Basal antennal article unarmed; outer Pereopods 2–4 (walking legs 1–3): Similar, slender, margin of article 2 with long slender spine, not reach- elongate; ornamentation similar in both sexes; seg- ing beyond article 4; article 3 unarmed; scaphocerite ments spinose, surface between major spines smooth or minute, blunt, shorter than article 4; article 4 unarmed, with few, scattered spines or granules. Pereopod 3 and about half as long as article 5. 4 subequal, longer than pereopod 2. Distal margins of Abdomen: Somite 1 with low triangular teeth on coxae unarmed; surface smooth. Ischiobasis with 3 or posterior margin. Somite 2 rugose; composed of 3 dis- 4 non-setose distal spines. Merus subcircular to ovate tinct plates separated by narrow groove; median sur- in cross section; extensor margin with 4–10 spines of face with pair of slender submedian spines; posterior which 2 or 3 are distinctly longer in addition to long, margin with 4–6 stout spines or angular prominences; prominent, distal spine; flexor margin with 2 rows of 4 lateral margins irregularly dentate. Somites 3–5 with or 5 small spines; upper surface with irregular row of nodular median plates; surface of submedian plates 5–8 small spines. Carpus about half merus length; ex- irregular, but unarmed; marginal plates of males sub- tensor margin with distal and second proximal spines divided, with angular or sharp apices; marginal plates longest, exceeding one-third carpus length; surface of females with left margin crenulate to spinose (fused with scattered small spines. Propodus ovate in cross with submedians), right marginals subdivided, with section; with 7–11 small spines on extensor margin angular to spiniform apices. Somite 6 of both sexes and 4–7 small spines on dorsal surface; flexor margin as long as or longer than wide, distal margin with 2 with 4–7 small spines. Dactylus curved, rounded in or 4 distal spines or prominences. Telson rounded, cross section distally; proximally with 2 spines on unarmed. either side adjacent to articulation; with 3 or 4 small Pereopod 1 (chelipeds): Dimorphic but spination simi- corneous spines along extensor margin; flexor margin lar. Surfaces of both chelipeds smooth or with scattered smooth or with about 4 soft, minute, widely spaced granules in addition to major spines. Coxa with tufts of setae; apex corneous. setae, unarmed; ischiobasis with 3 stout ventral spines Pereopod 2 length 2.91–3.21 pcl (male), 2.64–2.77 and about 6 blunt tubercles. Merus inner margin with pcl (female). Merus 1.10–1.16 pcl (male), 0.96 pcl stout subdistal spine and 2 or 3 smaller spines or acute (female);length:height ratio 6.56–7.42 (male), 6.43–8.64 tubercles; ventral margin with two rows of 2 or 3 low (female). Carpus 0.48–0.51 merus length (male), conical spines; dorsal and lateral surface spinose, spines 0.51–0.52 (female). Propodus 0.81–0.86 merus length largest distally. Carpus with prominent spines on dor- (male), 0.84–0.88 (female); length:height ratio 11.16– sal and lateral surfaces; dorsal margin with row of 3 12.8 (male), 10.09–13.35 (female). Dactylus 0.52–0.58 stout spines; ventral margin with small scattered acute propodus length (male), 0.62–0.63 (female). tubercles. Palms of both chelipeds in both sexes with Pereopod 3 length 3.12–3.48 pcl (male), 2.69–3.01 similar ornamentation; with small spines and acute pcl (female). Merus 1.10–1.23 pcl (male), 0.95–1.04 pcl tubercles on dorsal, lateral and ventral surfaces, inner (female); length:height ratio 6.00–7.44 (male), 6.46–6.94 surface with few low tubercles; dorsal margin with row (female). Carpus 0.51–0.53 merus length (male), 0.52 of 3 or 4 small conical spines; midlateral surface with 2 (female). Propodus 0.85–0.88 merus length (male), rows of 3 or 4 small spines or tubercles, of similar size 0.85–0.87 (female); length:height ratio 11.70–13.58 to dorsal row; ventral surface with scattered tubercles, (male), 10.67–11.06 (female). Dactylus 0.51–0.68 pro- smaller than lateral and dorsal spines. podus length (male), 0.57–0.61 (female). Major cheliped 1.38–1.59 pcl (male), 1.19–1.40 Pereopod 4 length 3.12–3.48 pcl (male), 2.57–2.95 (female); upper palm length 1.15–1.33 times height pcl (female). Merus 1.10–1.17 pcl (male), 0.92–1.02 pcl (male), 1.42–1.55 (female); occlusal margins of fingers (female); length:height ratio 6.88–7.84 (male), 6.59–7.33 corneous for distal third, proximally with 3 calcareous (female). Carpus 0.54–0.56 merus length (male), 0.54 nodules; dactylus dorsal margin broadly convex, with (female). Propodus 0.89–0.93 merus length (male), rows of tufts of golden setae and blunt proximal tooth, 0.88–0.90 (female); length:height ratio 12.32–13.57 1.22–1.31 times longer than dorsal margin of palm (male), 10.96–11.04 (female). Dactylus 0.50–0.56 pro- (male), 1.19–1.40 (female). podus length (male), 0.61–0.62 (female). Minor cheliped 1.38–1.50 pcl (male), 1.27–1.40 (female); upper palm length 1.35–1.49 times height Colour in life. Overall deep-red (Pl. 1C).

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Etymology. Named australiensis, for the Australian southern New South Wales (~35°30’S). The ranges of distribution of the new species. the two species overlap, and are sometimes sympatric off central and southern New South Wales. Differences Remarks. Lithodes longispina Sakai, 1971 was first between L. australiensis and L. richeri are outlined under reported from Australia based on “identification of the account of the latter. large numbers of L. longispina in south-east Austral- Lithodes australiensis sp. nov. is most similar to L. ian waters” from collections in the Australian Mu- robertsoni sp. nov. and L. aotearoa sp. nov. from New seum (Dawson & Yaldwyn 1985: 72). All records of Zealand, and L. rachelae sp. nov. from southwestern L. longispina from Australia and New Zealand, how- Australia, in having the combination of unarmed flexor ever, are referable to other species. As speculated by margins of the dactyli of the walking legs in adults, Macpherson (2001) and Poore (2004), many records of and long, slender, dorsal spines that persist in early L. longispina from New South Wales are based on L. adults. Although L. australiensis has in the past been richeri Macpherson, 1990, reported below. The second misidentified as L. longispina, the new species differs L. longispina-like species from southeastern Australia by its two long dorsal branchial spines: one anteriorly is herein named L. australiensis sp. nov. Lithodes austral- and one posteriorly. Lithodes longispina lacks the long iensis is a southern species, ranging from central New dorsal posterior branchial spine, at most bearing a short South Wales (~ 32°S) to the South Tasman Rise, south conical tooth or tubercle. Lithodes australiensis is distin- of Tasmania (~49°S). Lithodes richeri has a more north- guished from L. rachelae by its proportionally shorter erly distribution than L. australiensis, ranging from walking legs (see account of L. rachelae), and from L. Vanuatu and New Caledonia (possibly also Indonesia; robertsoni also by walking leg proportions (see account see account of L. richeri) to the vicinity of Brush Island, of L. robertsoni). Lithodes australiensis differs from L.

Figure 17. Geographic distribution of Lithodes australiensis sp. nov. (○), L. rachelae sp. nov. (●), and L. chaddertoni sp. nov. (?) Inset: bathymetric distribution of L. australiensis.

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ aotearoa in having 5 or 6 rather than 8–12 posterior present up to about 60 mm pcl, but the marginal plates branchial marginal spines and in having abdominal of abdominal somite 2 remain unfused to the median somite 2 composed of three plates in adults, rather than plate in adult males (adult females are presently un- being fused into a single plate as in adult L. aotearoa. known, but this feature is not known to be sexually Additionally, adult L. australiensis have more elongate dimorphic in other species of Lithodes). walking legs than adult L. aotearoa; in the former, the pereopod 4 merus length always exceeds the pcl Distribution. Southeastern Australia, from the vicinity (1.09–1.17 pcl), whereas in the latter, it only slightly of Cape Hawke, New South Wales, south to Victoria, exceeding pcl in the largest males (0.91–1.06) (see Table Tasmania and the South Tasman Rise; 540–1312 m, 1). Female L. australiensis also have more elongate walk- usually 1000–1100 m. ing legs than female L. aotearoa but the differences are less pronounced (Table 2). Lithodes australiensis possibly Lithodes chaddertoni sp. nov. matures at a larger size than does L. aotearoa because (Figs 17–20, Pl. 4A) the flexor spines on the dactyli of the walking legs are present up to about 60 mm pcl in L. australiensis, but Type material. Holotype: NMNZ Cr11193, male (cl 190.1 are lost by about 20 mm pcl in L. aotearoa. mm, pcl 100.9 mm, cw 89.9 mm), Naturaliste Plateau or All known female L. australiensis are juveniles, so the Broken Ridge, Indian Ocean off SW Western Australia, size at which females mature is not presently known, crab pot, coll. J. Chadderton, 2002. although it exceeds about 53.8 mm pcl (the largest known female). The abdomen of the 53.8 mm female Diagnosis. Carapace dorsal surface with long slender is already asymmetrical (with spinular and subdivided spines; gastric region with 4 upright spines, anterior right marginal plates, and fused left marginal and pair longer than half length of hepatic spine, posterior submedian plates), although it has not yet achieved pair half length of hepatic spine; cardiac and intestinal full asymmetry. regions each with 2 long upright spines; branchial sur- As in other lithodids, dorsal spine length in L. aus- face with 2 or 3 long upright spines. Rostrum exceeding traliensis becomes proportionally shorter with increas- 0.8 pcl. First and second primary marginal branchial ing body size. The longest carapace spine (hepatic) is spines greatly elongated, of similar length. Posterior as long as the pcl at 24.1 mm pcl (TM G3582), 0.7 pcl at branchial margin with 6 or 7 short spines, 4 prominent. 60.3 mm pcl (TM G3659), and 0.3 pcl in the 112.2 mm Chelipeds unequal; palms with low tubercles or spines pcl holotype. The row of corneous spines on the flexor on dorsal surface. Walking leg dactyli with spinous margin of the walking leg dactyli, a juvenile feature, is flexor margin; half propodi length.

Table 1. Selected morphometric measurements of adult male L. aotearoa sp. nov., L. australiensis sp. nov., L. rachelae sp. nov., L. robertsoni sp., and L. paulayi Macpherson & Chan, 2008. Measurements of L. paulayi based on male holotype, pcl 112.0 mm, cw 102.7 mm (FLMNH 2283).

Male pcl P4 L/pcl P4 merus L/pcl P4 merus L/H P4 propodus (Male) (Male) (Male) L/H (Male) L. aotearoa 72.3–194.0 2.68–2.98 0.91–1.06 5.56–7.16 9.12–13.04 L. australiensis 78.0–133.8 3.12–3.48 1.10–1.17 6.88–7.84 12.32–13.57 L. robertsoni 81.7–128.1 3.02–3.20 1.05–1.10 6.70–8.16 11.43–14.08 L. rachelae 75.0–113.1 3.62–3.71 1.29–1.30 9.17–9.36 15.24–15.61 L. paulayi 112.0 3.17 1.14 9.59 17.37

Table 2. Selected morphometric measurements of adult female L. aotearoa sp. nov., L. australiensis sp. nov., L. rachelae sp. nov., and L. robertsoni sp. nov. Females of L. paulayi not presently known.

Female pcl P4 L/pcl P4 merusL/pcl P4 merus L/H P4 propodus (Female) (Female) (Female) L/H (Female) L. aotearoa 69.6–161.3 .89–2.77 0.62–0.95 4.55–6.43 8.56–10.41 L. australiensis 53.8–84.5 2.57–2.95 0.92–1.02 6.59–7.33 10.96–11.04 L. robertsoni 70.2–71.2 2.91–2.92 0.96–0.98 6.11–6.76 11.55–11.87 L. rachelae 43.8 3.10 1.06 13.21 15.24–15.61 L. paulayi ? ? ? ? ?

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 18. Lithodes chaddertoni sp. nov., male holotype, cl 190.1 mm, pcl 100.9 mm, cw 89.9 mm, Western Australia (NMNZ Cr11193). A, dorsal habitus. B, carapace, right lateral view.

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 19. Lithodes chaddertoni sp. nov., male holotype, cl 190.1 mm, pcl 100.9 mm, cw 89.9 mm, Western Aus- tralia (NMNZ Cr11193). A, anterior carapace, dorsal view. B, right orbit and antenna. C, right chela. D, left chela. E, abdominal somite 2. F, right pereopod 3 dactylus.

Description. Carapace: Pyriform, about 1.13 times longer 0.9 pcl. Gastric region convex, with 2 pairs of long, than wide; regions indicated; dorsal surface armed upright spines. Cardiac region with pair of minute an- with long, slender spines and very few, scattered, terior spines and posterior pair of long upright spines, widely separated granules or small conical tubercles, posterior pair almost as long as hepatic spine. Hepatic surface otherwise smooth; longest spine (hepatic spine) spine directed anterolaterally. Posterior margin of

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ intestinal region with pair of long, upright spines. or acute tubercles. Palm with 7–11 small scattered Branchial surface with 2 long upright spines: first at spines or acute tubercles on dorsal and lateral surfaces, level of pereopod 3 coxa, second at level of pereopod inner and ventral surfaces unarmed. 4 coxa. Branchial margins spinose; anterior branchial Major cheliped 1.74 pcl; upper palm length 1.57 margin with 2 slender spines, posterior spine twice or times height; occlusal margins of fingers corneous for more length of anterior, half or more length of hepatic distal quarter, proximally with 3 calcareous nodules; spine; 1st primary marginal branchial spine as long as dactylus dorsal margin broadly convex, with rows of hepatic spine; lateral branchial margin with 1 or 2 short tufts of golden setae and blunt proximal tooth, 1.28 spines; 2nd primary marginal branchial spine slightly times longer than dorsal margin of palm. shorter than 1st primary marginal branchial spine; Minor cheliped 1.69 pcl; upper palm length 1.76 posterior branchial margin with 6 or 7 short spines, 4 times height; occlusal margin corneous for slightly less prominent. Pterygostomian region granulate, irregular, than distal half, proximally crenulate; dactylus dorsal with short subdistal tooth. margin broadly convex, with rows of tufts of golden Rostrum 0.88 pcl, comprising proximal and distal setae and blunt proximal tooth, 1.56 times longer than portions; proximal portion angled dorsally by about dorsal margin of palm. 45°, with pair of long, divergent dorsal spines that Pereopods 2–4 (walking legs 1–3): Similar, slender, overreach midlength of distal portion of rostrum; distal spinose; surface between major spines smooth or with portion subhorizontal, distally bifurcate for half length, few, minute, sparsely distributed spines. Pereopod 4 forming pair of strongly divergent spines; ventral ros- longest. Coxa of pereopods 2–3 unarmed; pereopod 4 tral spine elongate, extending anteriorly beyond apices coxa with anterodistal spine. Ischiobasis with 4 distal of outer orbital spine. Posterior orbital margin concave. spines, posteriormost spine of pereopod 4 longest. Outer orbital spine directed anteriorly, subequal to Merus subcylindrical; flexor margins with 5–8 spines; anterolateral spines, reaching beyond cornea. extensor margins with 2 or 3 prominent spines and Ocular peduncle: Longer than cornea; unarmed. 3–5 smaller spines, in addition to long, prominent, Antennule: Peduncle unarmed, reaching anteriorly distal spine, exceeding half merus length; dorsal sur- beyond antennal peduncle by three-quarters length of face with two irregular rows of 4–8 spines of which 3 distal antennular peduncle article. or 4 prominent. Carpus surface with scattered, short, Antenna: Basal antennal article unarmed; outer slender, strongly elongated distal and second proximal margin of article 2 with slender spine that reaches spines, each exceeding carpus length. Propodus with to or slightly beyond article 4; articles 3–5 unarmed; 8–11 spines on extensor and flexor margins. Dacty- scaphocerite short, blunt; article 4 about half as long as lus curved, laterally compressed; proximally with 2 article 5. Peduncle extending beyond apex of ventral spines on either side adjacent to articulation; with 4–6 rostral spine by two-thirds length of distal peduncular slender corneous spines along extensor margin; flexor article. margin lined with 7–10 slender corneous spines; apex Abdomen: Somite 1 with pair of tubercles or trian- corneous. gular teeth on posterior margin. Somite 2 composed of Pereopod 2 length 3.12 pcl. Merus 1.12 pcl; length: three plates; surface sparsely granulate; median sur- height ratio 8.79. Carpus 0.51 merus length. Propodus face with pair of slender submedian spines; posterior 0.87 merus length; length:height ratio 14.80. Dactylus margin with 8 spines of varying length; lateral margins 0.52 propodus length. irregularly dentate. Somites 3–5 with median nodules; Pereopod 3 length 3.39 pcl. Merus 1.16 pcl; length: submedian plates pitted, unarmed; marginal plates height ratio 8.74. Carpus 0.54 merus length. Propodus with irregularly angular margins. Somite 6 elongate, 0.91 merus length; length:height ratio 15.17. Dactylus unarmed, distal margin with 4 low, blunt prominences. 0.52 propodus length. Telson unarmed, rounded. Pereopod 4 length 3.5 pcl (estimated; dactylus bro- Pereopod 1 (chelipeds): Dimorphic but spination ken). Merus 1.15 pcl; length:height ratio 8.80. Carpus similar. Surfaces of both chelipeds smooth or with 0.56 merus length. Propodus 0.92 merus length; length: scattered granules in addition to several small spines. height ratio 14.84. Coxa unarmed; with tufts of setae mesially. Ischioba- sis with 4 stout ventral spines and about 3 or 4 blunt Colour in life. Overall deep-red (Pl. 4A). tubercles. Merus inner margin, with curved subdistal spine; ventral margin with 3 low conical spines and Etymology. Named in honour of John Chadderton, who scattered tubercles; dorsal margin with 5–7 small, collected the holotype. scattered spines and 3 prominent spines, increasing in length distally, distalmost exceeding half carpus Remarks. Lithodes chaddertoni sp. nov. is the most length; lateral surface spinose, spines largest distally. strongly spinose of known species of the genus, with Carpus with 6 spines and 3 or 4 scattered, small spines spines measuring up to 0.87 pcl, and rostral length

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 20. Lithodes chaddertoni sp. nov., male holotype, cl 190.1 mm, pcl 100.9 mm, cw 89.9 mm, Western Australia (NMNZ Cr11193). A, carapace. B, abdomen.

0.88 pcl in adults. Lithodes chaddertoni resembles Distribution. Presently known only from off southwest- L. jessica sp. nov. from New Zealand in the combina- ern Western Australia. tion of long dorsal spines in adults and spinose flexor margins of the walking leg dactyli. The two species Lithodes jessica sp. nov. (Figs 21–26) are readily distinguished by cheliped form: equal with smooth palms in L. jessica versus distinctly dimorphic Type material. Holotype: NIWA 61195, ovigerous female with spinose palms in L. chaddertoni. Additionally, (cl 97.8 mm, pcl 58.6 mm, cw 52.1 mm), southern Lord L. chaddertoni has very few carapace tubercles and Howe Rise, 35°56.1’S, 166°09.7’E, 1050 m, trip 1204/27, granules compared to L. jessica, and proportionally Z9720, coll. A. Knox, 6 Mar 1999. longer major spines, despite the large size difference Paratypes: NIWA 34902, 1 ovigerous female (cl 110.5 (maximum 100.9 mm in L. chaddertoni versus 64.1 mm mm, pcl 64.1 mm, cw 58.1 mm), Lord Howe Rise, no other pcl in L. jessica) (compare Figs 18 and 21). data; NMNZ Cr11138, 1 male (cl 70+ mm, pcl 48.5 mm, Many species of Lithodes have long slender dorsal cw 43.7 mm), off Napier, Hawkes Bay, 39°59’S, 178°13’E, spines in juveniles, but spine length is usually much 680–1100 m, FV Peterson, coll. S. O’Shea, Apr 1994. reduced in adults. Thus, adult L. chaddertoni and L. jessica are apparently paedomorphic, closely resem- Diagnosis. Carapace dorsal surface with long slender bling juvenile L. australiensis sp. nov. and L. aotearoa sp. spines; gastric region with 4 spines, anterior pair nov. in the long dorsal spines and armed flexor margins greatly elongate, subequal to length of hepatic spine, of the dactyli of the walking legs. posterior pair short, less than one-quarter length of Adults of other ‘long-spined’ species of Lithodes anterior pair; cardiac and intestinal regions each with 2 from southern Australia, namely L. australiensis and long upright spines; branchial surface with 2 or 3 long L. rachelae, lack a spinose flexor margin on the walk- upright spines. Rostrum about 0.7 pcl. First and second ing leg dactyli and have proportionally shorter dorsal primary marginal branchial spines short, less than spines. Note that flexor dactylar spines are also present one-third length of hepatic spine. Posterior branchial in juvenile L. australiensis up to about 60 mm pcl; juve- margin with 7 or 8 spines. Chelipeds equal, slender; niles of L. rachelae are presently unknown.

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 21. Lithodes jessica sp. nov., ovigerous female holotype, cl 97.8 mm, pcl 58.6 mm, cw 52.1 mm, Lord Howe Rise (NIWA 61195). A, dorsal habitus. B, carapace, right lateral view. 42

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 22. Lithodes jessica sp. nov., ovigerous female holotype, cl 97.8 mm, pcl 58.6 mm, cw 52.1 mm, Lord Howe Rise (NIWA 61195). A, right cheliped. B, right orbit and antenna. C, left pereopod 4. D, left pereopod 4 dactylus. E, posterior carapace and abdominal somite 2. F, abdomen.

palms cylindrical, smooth, with very few low tubercles Description. Carapace: Pyriform, about 1.10–1.12 times or short teeth. Walking leg dactyli half propodi length; longer than wide; regions indicated; dorsal surface dactyli with spinous flexor margin. armed with long, slender spines and numerous, well- spaced granules or small conical tubercles, surface

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 23. Lithodes jessica sp. nov., ovigerous female holotype, cl 97.8 mm, pcl 58.6 mm, cw 52.1 mm, Lord Howe Rise (NIWA 61195). A–B, carapace.

otherwise smooth. Gastric region convex, with 1 pair vergent spines; ventral rostral spine elongate, extend- of long, upright spines anteriorly, posteriorly with ing anteriorly beyond apices of outer orbital spine. pair of short spines or acute tubercles; anterior gastric Posterior orbital margin rounded; outer orbital spine spines longest of carapace spines (0.6–1.0 pcl). Cardiac directed anteriorly, shorter than first hepatic spine, not region with pair of anterior tubercles or small spines reaching beyond cornea. and posterior pair of long upright spines. Hepatic spine Ocular peduncle: Longer than cornea; unarmed. directed anterolaterally. Posterior margin of intestinal Antennule: Peduncle unarmed, reaching anteriorly region with pair of long, upright spines. Branchial sur- beyond antennal peduncle by half to three-quarters face with 2 long upright spines: first at level of anterior length of distal antennular peduncle article. margin of pereopod 4 coxa, second at level of posterior Antenna: Basal antennal article unarmed; outer margin of pereopod 4 coxa, often with short spine at margin of article 2 with slender spine that reaches to base. Branchial margins spinose; anterior branchial or slightly beyond midlength of article 4; articles 3–5 margin with 2 short spines, posterior spine the longer; unarmed; scaphocerite absent; article 4 about half as 1st primary marginal branchial spine one-quarter length long as article 5. Peduncle extending beyond apex of hepatic spine; lateral branchial margin with 3 short ventral rostral spine by half to three-quarters length of spines; 2nd primary marginal branchial spine about distal peduncular article. one-eighth length of hepatic spine; posterior branchial Abdomen: Somite 1 with pair of tubercles or trian- margin with 7 or 8 spines. Pterygostomian region gular teeth on posterior margin. Somite 2 undivided, granulate or tuberculate, anterior margin angular. surface sparsely granulate, submedian plates fused Rostrum 0.67–0.72 pcl, comprising proximal and with median in adults; median surface with pair of distal portions; proximal portion angled dorsally by stout submedian spines; posterior margin with 4 stout about 45°, with pair of long, divergent dorsal spines spines or angular prominences; lateral margins irregu- that overreach midlength of distal portion of rostrum; larly dentate. Somites 3–5 with nodular median plates; distal portion subhorizontal, distally bifurcate for submedian plates pitted, unarmed; marginal plates about two-thirds length, forming pair of strongly di- with irregularly angular margins in males; marginal

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 24. Lithodes jessica sp. nov., male paratype, pcl 48.5 mm, cw 43.7 mm, off Napier (NMNZ Cr11138). A, dorsal habitus. B, carapace. C, abdomen. 45

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 25. Lithodes jessica sp. nov., male paratype, pcl 48.5 mm, cw 43.7 mm, off Napier (NMNZ Cr11138). A, right cheliped. B, right cheliped. C, left pereopod 4. D, left pereopod 4 dactylus.

plates of females with left margin smooth to crenulate calcareous, crenulate; dactylus dorsal margin straight (fused with submedians), right marginals subdivided for proximal two-thirds, 1.06–1.08 times longer than into 3 or 4 plates on right side, with spiniform apices. dorsal margin of palm (male), 1.12 (female). Somite 6 of both sexes elongate, unarmed, distal mar- Pereopods 2–4 (walking legs 1–3): Similar, slender, gin with 4 low, blunt prominences. Telson unarmed, spinose; surface between major spines smooth or with rounded. few, minute, sparsely distributed spines. Pereopod 4 Pereopod 1 (chelipeds): Subcylindrical; slender, sub- longest. Coxae unarmed. Ischiobasis with 2 distal ven- equal, dimorphism not marked. Chelipeds 1.83 pcl tral spines, posteriormost spine of pereopod 4 longest. (male), 1.55–1.60 pcl (female). Coxa unarmed; mesial Merus subcylindrical; extensor and flexor margins with surfaces with setal tufts; ischiobasis with 3 stout ven- 2–4 longer spines amongst small, scattered spinules, re- tral spines. Merus inner margin smooth or granular; spectively in addition to long, prominent, distal spine; extensor margin with slender distal spine and 7–9 small dorsal surface with 5–7 spines and scattered spinules. spines or spinules; flexor margin with 1 or 2 spinules. Carpus surface with small scattered spines; extensor Carpus longer than palm; smooth except for 4 promi- margin with distal and second proximal spines longest, nent spines on proximal half and 2–4 scattered spinules exceeding three-quarters carpus length. Propodus with distally. Palm smooth, unarmed; upper palm length 7–9 spines on extensor and flexor margins. Dactylus 2.95–3.00 times height (male), 2.72 (female). Fingers half propodus length; curved, laterally compressed; unarmed; with rows of tufts of golden setae; occlusal proximally with 2 spines on either side adjacent to ar- margins corneous for distal third to half, remainder ticulation; extensor margin with 3–5 slender corneous

spines along proximal three-quarters; length of flexor length:height ratio 9.28 (male), 7.14–8.21 (female). margin lined with 8 or 9 slender corneous spines; apex Carpus 0.51 merus length (male), 0.59–0.61 (female). corneous. Propodus 0.83 merus length (male), 0.96 pcl (female); Pereopod 2 length 3.45 pcl (male), 2.81–2.95 pcl length:height ratio 15.50 (male), 14.02–14.14 (female). (female). Merus 1.25 pcl (male), 0.94–1.02 pcl (female); Dactylus 0.52 propodus (male), 0.49–0.51 (female). length:height ratio 9.01 (male), 7.71–8.48 (female). Pereopod 4 length 3.54 pcl (male), 2.99–3.12 pcl Carpus 0.50 merus length (male), 0.57–0.60 (female). (female). Merus 1.34 pcl (male), 0.91–1.01 pcl (female); Propodus 0.84 merus length (male), 0.90–0.93 (female); length:height ratio 9.69 (male), 6.44–7.84 pcl (female). length:height ratio 15.55 (male), 14.02–14.69 (female). Carpus 0.49 merus length (male), 0.62–0.67 (female). Dactylus 0.50 propodus length (male), 0.51–0.53 (fe- Propodus 0.83 merus length (male), 1.02–1.04 (female); male). length:height ratio 16.64 (male), 14.11–15.08 (female). Pereopod 3 length 3.42 pcl (male), 2.98–3.11 pcl Dactylus 0.52 propodus (male), 0.49–0.54 (female). (female). Merus 1.26 pcl (male), 0.98–1.04 pcl (female);

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Colour in life. Not known. mm, cw 88.3 mm), off Garden Cove, Macquarie Island, 54°30’S, 158°57’E, 16 m, 20 Nov 1977; NIWA 61160, Etymology. Named for Jessica Ahyong; used as a noun in 1 juvenile female (cl 31.3 mm, pcl 20.9 mm, cw 18.5 apposition. mm), Macquarie Ridge, S of Macquarie Island, 55°00’S, 158°46.99’E, 110 m, C730, 24 Nov 1961; NIWA 61187, Remarks. Owing to the long dorsal spines, includ- 1 male (cl 136.5 mm, pcl 111.1 mm, cw 106.0 mm), ing two long dorsal branchial spines, Lithodes jessica Macquarie Island, 54°29.5’S, 158°58.50’E, 22 m, C732, sp. nov. resembles early adults of L. aotearoa sp. nov. 25 Nov 1961; NIWA 61186, 1 male (cl 90.9 mm, pcl 73.1 (pcl ~100 mm or less) and L. robertsoni sp. nov. from mm, cw 71.3 mm), Macquarie Ridge, N of Macquarie New Zealand, L. chaddertoni sp. nov., L. rachelae sp. Island, 54°25.00’S, 159°01.99’E, 104 m, C733, 25 Nov nov., and L. australiensis sp. nov. from Australia, 1961; NIWA 40798, 1 male (pcl 99.3 mm, cw 94.1 mm), L. megacantha Macpherson, 1991 from French Polynesia, Macquarie Ridge, 55°21.19–21.36’S, 158°26.20–26.46’E, and L. paulayi Macpherson & Chan, 2008, from Guam. 605–709 m, TAN0803/093 SEL 1483, 15 Apr 2008; Lithodes jessica is most obviously distinguished from NMV J14503, 1 female (pcl 68.3 mm, cw 62.6 mm), each of these species by the subequal, slender, cylin- Garden Cove, Macquarie Island, coll. K. Bennett, 2 drical chelipeds, which bear only a few short spines Feb 1976; NMV J59364, 2 males (cl 121.5–133.0 mm, on the merus and carpus, and none on the palm. The pcl 100.6–113.2 mm, cw 98.3–113.5 mm), 54°35.42’S, chelipeds of the aforementioned species are instead 158°56.60–58.42’E, 250–500 m, SS0199/93, 26 Jan 1999; multispinose and distinctly dimorphic, in which NMV J14550, 1 male (cl 145.1 mm, pcl 120.0 mm, cw the major palm is highest near the dactylar articula- 116.8 mm), Camp Beach, Macquarie Island, coll. P. tion. Lithodes jessica also differs from L. australiensis, Puckert, 3 June 1957; SAM C6852, 1 juvenile male (cl L. aotearoa, L. rachelae, L. megacantha, L. paulayi, and 27.8 mm, pcl 16.6 mm, cw 14.8 mm), W side of Mac- L. robertsoni by the spinose versus unarmed flexor quarie Island, 54°42.7–41.9’S, 158°46.1–45.9’E, 100–300 margin on the walking leg dactyli in adults, and by the m, dredge, RV Southern Surveyor, SS0199/60, 22 Jan longer walking legs. In this respect, L. jessica resembles 1999. L. chaddertoni, which also has a spinular flexor margin of the walking leg dactyli. Note, however, that the Other material examined. Campbell Plateau: NIWA flexor spination of the walking leg dactyli, a juvenile 34897, 1 damaged juvenile male (pcl 44.4 mm, cw 40.1 lithodid feature retained by adults of L. jessica and mm), W of Auckland Islands, 50°11’S, 165°49.3’E, 1090 L. chaddertoni, is also present in juvenile L. australiensis m, MFISH trip 1292/5, FV Atlantic Elizabeth, Z10222, and L. aotearoa up to about 60 mm pcl and 20 mm pcl, coll. S. Beatson, 18 Nov 1999. respectively. Therefore, walking leg dactyl armature Solander Trough: NIWA 46529, 1 juvenile (cl 16.1 should not be used to separate specimens of L. jessica mm, pcl 7.9 mm cw 6.6 mm), 50°56.31’S, 164°33.16’E, from small L. australiensis and L. aotearoa. Instead, the 1053–998 m, TAN0306/4, 14 Apr 2003; NIWA 46524, 1 cheliped dimorphism, barely evident in L. jessica, is juvenile female (cl 24.9 mm, pcl 12.6 mm, cw 10.9 mm), obvious even in the smallest specimens of L. aotearoa 50°56.56’S, 164°36.55’E, 1140–1105 m, TAN0306/6, 14 and L. australiensis examined. Apr 2003.

Distribution. Presently known only from northern New Material studied from photographs. Macquarie Ridge: Zealand waters between the southern Lord Howe Rise NIWA 40735, 1 juvenile male (pcl 35.9 mm), 55°21.69– and Hawkes Bay; 680–1100 m. 21.73’S, 158°25.66–26.11’E, 501–630 m, TAN0803/91, 15 Apr 2008; NIWA 40902, 1 juvenile female (pcl 43.8 Lithodes macquariae sp. nov. mm), 56°14.77–14.49’S, 158°30.33–30.90’E, 676–750 m, (Figs 27–31, Pl. 2E) TAN0803/98, 16 Apr 2008.

Lithodes murrayi. — Hale, 1941: 272, pl. 3: figs. 3, 4. — Healy & Diagnosis. Carapace dorsal spines short and conical, Yaldwyn, 1970: 74, fig. 37. —Dawson, 1988: 225. — Davie, with 4 gastrics, 2 cardiacs, 2 branchials and 2 intesti- 2002: 73 [Macquarie Island occurrences]. — Zaklan, nals. Hepatic margin short tubercle or spine behind 2002: 766 [Macquarie Island occurrences] [Not L. murrayi anterolateral spine. Posterior branchial margin with Henderson, 1888]. 6–9 triangular spines. Rostrum about 0.2–0.3 pcl, with pair of short divergent dorsal spines; distal portion dis- Type material. Holotype: AM P42530, male (cl 123.2 tally bifurcate for one-third length. Abdominal somite mm, pcl 102.0 mm, cw 108.3 mm), off Garden Cove, 2 composed of three segments, with marginal plates Macquarie Island, 54°30’S, 158°57’E, 16 m, MA-7, rocky clearly demarcated. Chelipeds dimorphic, segments bottom, coll. R. Williams, 20 Nov 1977. spinose. Walking legs with surface between major Paratypes: AM P42531, 1 male (cl 107.5 mm, pcl 89.3 spines smooth or with scattered spines or granules;

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 27. Lithodes macquariae sp. nov., male holotype, cl 123.2 mm, pcl 102.0 mm, cw 108.3 mm, off Garden Cove, Macquarie Island (AM P42530). A, dorsal habitus. B, carapace. C, right pereopod 4. 49

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 28. Lithodes macquariae sp. nov., male holotype, cl 123.2 mm, pcl 102.0 mm, cw 108.3 mm, off Garden Cove, Macquarie Island (AM P42530). A, right chela. B, left chela. C, posterior carapace and abdominal somite 2. D, ventral surface and abdomen. E, carapace, right lateral view. F, right orbit and antenna.

longest dorsal spines not exceeding half merus height. Description. Carapace: Pyriform, 0.94–1.09 times longer Walking leg 2 longest, about 3 times pcl in males; merus than wide; regions indicated; dorsal surface armed as long as pcl in males, shorter than pcl in females, with short, conical spines and scattered, widely sepa- less than 6 times longer than high; propodus shorter rated granules. Gastric region convex, with 2 pairs of than merus; dactylus about 0.6 propodus length, flexor short, conical spines. Cardiac region with pair of low margin unarmed. blunt anterior tubercles and posterior pair of short coni-

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 29. Lithodes macquariae sp. nov., female paratype, pcl 68.3 mm, cw 62.6 mm, Garden Cove, Macquarie Island (NMV J14503). A, dorsal habitus. B, abdomen. 51

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ cal spines. Hepatic margin with 1 or 2 short tubercles with 3 stout ventral spines and about 6 small tubercles. or small spines near midlength and anterolaterally Merus inner margin, with stout subdistal spine and 2 directed conical spine near junction with branchial or 3 smaller spines; ventral surface with two rows of 2 region. Intestinal region with pair of short, upright, or 3 low tubercles; dorsal and lateral surface spinose, conical spines on posterior margin. Branchial surface spines largest distally. Carpus with prominent spines with 1 short spine at level of pereopod 3 coxa, 1 short on dorsal and lateral surfaces; dorsal margin with row spine at level of pereopod 4 coxa, and conical tuber- of 4 stout spines; lateral surface with 3 stout spines cle behind base. Branchial margins spinose; anterior of similar size to dorsal row in addition to scattered branchial margin with 2 short spines, posterior spine smaller spines. Palms of both chelipeds in both sexes the longer; 1st primary marginal branchial spine sub- with similar ornamentation; with small spines and equal to shorter than hepatic spine; lateral branchial acute tubercles on dorsal, lateral and ventral surfaces, margin with 2 or 3 short spines; 2nd primary marginal inner surface with acute tubercles; dorsal margin branchial spine shorter than 1st primary marginal with row of about 4 conical spines; midlateral surface branchial spine; posterior branchial margin with 6–9 with 2 rows of 4–7 small spines; ventral surface with spines. Pterygostomian region granulate or tubercu- 2 rows of short spines or tubercles (outer with 5 or 6, late, with small, anterior, submarginal spine. inner with 2). Rostrum short, 0.17–0.24 pcl, comprising upraised Major cheliped 1.51–1.60 pcl (male), 1.22 (female); proximal portion and horizontal distal portion; proxi- upper palm length 1.07–1.22 times height (male), 1.28 mal portion angled dorsally by at least 30°, with pair (female); occlusal margins of fingers corneous for distal of short, divergent dorsal spines; distal portion distally third, proximally with 3 calcareous nodules; dactylus bifurcate for one-third length; ventral rostral spine dorsal margin broadly convex, with rows of tufts stout, extending anteriorly beyond apices of outer of golden setae and blunt proximal tooth, 1.14–1.25 orbital spine but not beyond distal antennal peduncle times longer than dorsal margin of palm (male), 1.30 segement 4. Posterior orbital margin concave; outer (female). orbital spine directed anteriorly, not reaching beyond Minor cheliped 1.40–1.43 pcl (male), 1.22 (female); cornea; longer than anterolateral spine. upper palm length 1.17–1.35 times height (male), 1.29 Ocular peduncle: Longer than cornea, unarmed. (female); occlusal margin corneous for slightly less Antennule: Peduncle unarmed, reaching anteriorly than distal half, proximally crenulate; dactylus dorsal beyond antennal peduncle by about three-quarters margin broadly convex, with rows of tufts of golden length of distal antennular peduncle article. setae and blunt proximal tooth, 1.43–1.58 times longer Antenna: Basal antennal article unarmed; outer than dorsal margin of palm (male), 1.45 (female). margin of article 2 with slender spine, not reaching Pereopods 2–4 (walking legs 1–3): Similar, slender, beyond midlength of article 4; articles 3–5 unarmed; elongate; segments spinose, surface between major scaphocerite minute, shorter than article 4; apex blunt, spines smooth or with scattered spines or granules. or rarely acute; article 4 unarmed, about half as long Pereopod 4 longest, about 3 times pcl in males. Distal as article 5. margins of coxae with 2 short spines. Ischiobasis with Abdomen: Somite 1 with pair of triangular teeth on 3 non-setose distal spines. Merus subcircular to ovate posterior margin. Somite 2 composed of three plates, in cross section; extensor margin with 5–8 spines of with marginal plates clearly demarcated; surface which 3 or 4 are distinctly longer in addition to paired sparsely granulate; median surface with pair of stout distal spines; flexor margin with 2 rows of 4–7 spines; submedian spines; posterior margin with 4 posterior upper surface with row of 5–7 spines in addition to spines and several small acute tubercles; lateral mar- scattered smaller spines. Carpus slightly longer than gins irregularly dentate. Somites 3–5 with nodular half merus length; extensor margin with distal and median plates; surface of submedian plates irregular, second proximal spines longest; surface with scattered with scattered, low, acute tubercles; marginal plates small spines. Propodus ovate in cross section, shorter of males subdivided, with angular or sharp apices; than merus; with 6–9 spines on extensor margin and marginal plates of females with left margin crenulate scattered spines on dorsal surface; flexor margin with to spinose (fused with submedians), right marginals 5 or 6 smaller spines. Dactylus curved, rounded in subdivided, with angular to spiniform apices. Somite cross section distally; proximally with 2 spines on 6 of both sexes as long as or longer than wide, with either side adjacent to articulation; with 2–4 corneous 2 proximal 4 or distal prominences. Telson rounded, spines along extensor margin; flexor margin unarmed; unarmed, at most with 4 blunt granules. apex corneous. Pereopod 1 (chelipeds): Dimorphic but spination Pereopod 2 length 2.58–2.73 pcl (male), 2.17 (fe- similar. Surfaces of both chelipeds sparsely covered male). Merus 0.92–0.95 pcl (male), 0.75 (female); length: with granules in addition to major spines. Coxa with height ratio 4.48–5.33 (male), 4.73 (female). Carpus blunt tubercles and tufts of setae, unarmed; ischiobasis 0.53–0.55 merus length (male), 0.58 (female). Propodus

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 30. Lithodes macquariae sp. nov., A–B, juvenile male paratype, cl 27.8 mm, pcl 16.6 mm, cw 14.8 mm, W side of Macquarie Island (SAM C6852). C, female paratype, cl 31.3 mm, pcl 20.9 mm, cw 18.5 mm, Macquarie Ridge, S of Macquarie Island (NIWA 61160). 0.85–0.86 merus length (male), 0.88 (female); length: height ratio 5.20–5.92 (male), 5.05 (female). Carpus height ratio 7.55–8.80 (male), 8.41 (female). Dactylus 0.52–0.55 merus length (male), 0.57 (female). Propodus 0.59–0.67 propodus length (male), 0.59 (female). 0.84–0.90 merus length (male), 0.87 (female); length: Pereopod 3 length 2.72–2.99 pcl (male), 2.37 (fe- height ratio 8.23–9.05 (male), 8.35 (female). Dactylus male). Merus 0.96–1.05 pcl (male), 0.82 (female); length: 0.57–0.63 propodus length (male), 0.57 (female).

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Pereopod 4 length 2.82–2.98 pcl (male), 2.48 (fe- Remarks. Lithodes macquariae sp. nov. most closely male). Merus 0.95–1.00 pcl (male), 0.81 (female); length: resembles L. murrayi Henderson, 1888, with which it height ratio 4.76–5.90 (male), 5.21 (female). Carpus has been previously misidentified (Hale 1941; Healy 0.52–0.56 merus length (male), 0.59 (female). Propodus & Yaldwyn 1970). The Macquarie Island and south- 0.90–0.92 merus length (male), 0.93 (female); length: west Indian Ocean forms are no doubt closely related, height ratio 7.91–9.76 (male), 8.98 (female). Dactylus but differ consistently, justifying specific separation. 0.58–0.64 propodus length (male), 0.57 (female). Lithodes macquariae differs from L. murrayi chiefly in the proportional lengths of the walking leg dactyli, Colour in life. Deep-red overall. Juveniles usually measuring about 0.6 rather than less than 0.5 propodus paler, being pinkish overall with red spines, rostrum length, and in bearing 6–9 triangular posterior branchial and cephalic appendages (Pl. 1F). marginal spines (of which 6 or 7 are prominent) rather than 4–6 (of which 3 or 4 are prominent). Adults might Etymology. The specific name alludes to the distribu- also have a proportionally shorter rostrum, measuring tion of the species, the Macquarie Ridge; used as a noun about 0.2 pcl versus about 0.3 pcl in adults of 89 mm in the genitive singular. pcl or greater (Macpherson 1988c). Macpherson (1988c)

Figure 31. Geographic distributions of Lithodes macquariae sp. nov. (○) and L. robertsoni sp. nov. (●).

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ noted that rostral length should be used with caution known only from the southwestern Indian Ocean; in view of allometric variation and anomalies caused recent reports of L. murrayi from Peter I Island and the by damage and regeneration. However, morphometric Bellingshausen Sea (Klages et al. 1995; García Raso et data based on extensive series of L. murrayi from the al. 2005) require verification. Crozet Islands indicates that the rostral length follows a Of all lithodids known from the region, consistent pattern (Arnaud & Do-Chi 1977), so the dif- L. macquariae is the only species to range into shal- ferences observed in proportional rostral length might low sublittoral depths. Lithodes macquariae is known be real rather than apparent. Data from a larger series from 16–709 m depth around Macquarie Island, and of L. macquariae are required to better evaluate rostral 998–1050 m at 5° further north, near the Auckland length variation. Note that Henderson’s (1888) original Islands. This is consistent with the polar emergence figure ofL. murrayi incorrectly depicts the walking leg pattern observed in other lithodids at high latitudes, dactyli as about 0.6 propodus length. Macpherson’s particularly in the southern hemisphere (Lovrich et (1988b) redescription and figures of L. murrayi show al. 2002). Lithodes macquariae is one of two lithodid that the dactyli measure half or slightly less than half species known from the Macquarie Ridge. The second propodus length. species, Paralomis birsteini, can be distinguished by its The smallest juveniles examined (male pcl 7.9 mm, undivided abdominal somite 2, which, in L. macquariae, female pcl 12.6 mm, male pcl 16.6 mm, female pcl 20.9 comprises three plates. mm), differ from adults in expected features: a proportionally longer rostrum (1.0 pcl, 1.0 pcl, 0.7 pcl, and Distribution. Known only from the Macquarie Ridge 0.5 pcl, respectively) with longer apical spines, a more near Macquarie Island, north to the Solander Trough elongate carapace (pcl 1.2, 1.2, 1.1, 1.1 times cw, re- in the vicinity of the Auckland Islands; 16–1140 m. spectively), proportionally longer carapace spines, and flexor spinules on the walking leg dactyli. The length of Lithodes rachelae sp. nov. the hepatic spine is about 1.0 pcl at 7.9 and 12.6 mm pcl, (Figs 17, 32–34, Pl. 1D) and 0.8 pcl at 16.6 mm pcl. At 20.9 mm pcl, the hepatic spine is 0.3 pcl, which is considerably shorter than that ?Lithodes aff. longispina. — Poore et al., 2008: 27. of size-matched juveniles, or even juveniles of twice the size of L. australiensis in which the hepatic spine Type material. Holotype: SAM C6357, male (cl 142.0 mm, length exceeds the pcl. In the damaged male juvenile pcl 82.9 mm, cw 75.0 mm), Great Australian Bight, ap- (pcl 44.4 mm, NIWA 34897), the flexor spinules on the prox. 120 nautical miles [222 km] SSE of Eucla, South walking leg dactyli are minute and barely evident. Two Australia, 33°37.2’S, 129°52.0’E, 930–1087 m, FV Longva juvenile L. macquariae, a male (pcl 35.9 mm) and female III, shot 31, coll. K. Gowlett-Holmes, 5 Apr 1989. (pcl 43.8 mm), were captured and photographed in Paratypes: SAM 6853, 1 female (cl 80.0 mm, pcl 43.8 2008 by NIWA expedition TAN0803 to the Macquarie mm, cw 40.0 mm), SE of Albany, Western Australia, Ridge (Pl. 1F). Unfortunately, both specimens were lost 1500 m, FV Comet, coll. D. Turner, 10 Nov 1990; SAM at sea, but the relatively short carapace and pereopod C6367, 1 male (pcl 117.3 mm, cw113.1 mm), Great Aus- spines are evident in the photographs. tralian Bight, approx. 120 nautical miles SSE [222 km] Sexual dimorphism in L. macquariae is typical of of Eucla, South Australia, 33°45’S, 129°17’E, 999–1101 other lithodids. Adult females have an asymmetrical m, FV Adelaide Pearl, coll. K. Gowlett-Holmes & H. abdomen and proportionally shorter walking legs. The Cameron, 1 Aug 1988. examined female specimen is mature, having a fully developed abdomen and setose ventral surfaces of the Other material examined. Western Australia: NMV pereopodal coxae. J54985, 1 juvenile female (pcl 6.7 mm, cw 5.4 mm), SE Prior to revision by Macpherson (1988b, c), Lithodes of Albany, 35°26.05’S, 118°21.07’E, SS1005/028, RV murrayi was attributed a southern circumpolar distri- Southern Surveyor, 23 Nov 2005. bution, having been reported from the southwestern Indian Ocean (Prince Edward Island, Crozet Islands, Diagnosis. Carapace dorsal surface with slender Reunion Islands, South Africa, Macquarie Island spines; gastric region with 4 upright spines, anterior and New Zealand) (Yaldwyn & Dawson 1970). As pair longer than half length of hepatic spine, posterior shown by Macpherson (1988b, c), however, records pair half length of hepatic spine; cardiac and intestinal of L. murrayi from South Africa are referable to regions each with 2 long upright spines; branchial L. mamillifer Macpherson, 1988c, and those from Chile surface with 2 long upright spines. First primary are based on L. turkayi Macpherson, 1988b. Similarly, marginal branchial spine about two-thirds length of records of L. murrayi from the Australian and New hepatic spine. Branchial margins spinose; posterior Zealand regions are referable to L. macquariae and branchial margin with 7 or 8 short spines. Rostrum L. aotearoa sp. nov. Lithodes murrayi is thus reliably exceeding 0.6 pcl; proximal half strongly upraised by

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 32. Lithodes rachelae sp. nov., male holotype, cl 142.0 mm, pcl 82.9 mm, cw 75.0 mm, Great Australian Bight, SSE of Eucla, South Australia (SAM C6357). A, dorsal habitus. B, carapace, right lateral view.

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 33. Lithodes rachelae sp. nov., male holotype, cl 142.0 mm, pcl 82.9 mm, cw 75.0 mm, Great Australian Bight, SSE of Eucla, South Australia (SAM C6357). A, right chela. B, left chela. C, posterior carapace and abdominal somite 2. D, ventral surface and abdomen. E, carapace. F, right pereopod 4.

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ about 30°. Abdominal somite 2 comprising 3 plates in posterior margin. Somite 2 rugose; composed of 3 juveniles through adults. Antennal peduncle article 2 distinct plates separated by narrow groove; median with slender outer spine not reaching beyond article surface with pair of slender submedian spines; poste- 4. Chelipeds unequal; palms with low tubercles or rior margin with 6 stout spines; lateral margins irregu- spines on dorsal surface. Walking legs spinose; sur- larly dentate. Somites 3–5 with nodular median plates; face between major spines smooth or with very few, surface of submedian plates irregular, unarmed; mar- scattered spines. Pereopod 4 merus about 1.3 pcl in ginal plates of males subdivided, with angular apices; males, as long as pcl in females; merus length about 9 marginal plates of females with left margin crenulate times height (male), less than 8 times height (female); (fused with submedians), right marginals subdivided, propodus length 15–16 times height (male), 13 times with angular apices. Somite 6 of both sexes longer than height (female); dactylus flexor margin unarmed. wide, distal margin with 2 or 4 blunt prominences. Telson rounded, unarmed. Description. Carapace: Pyriform, 1.04–1.10 times longer Pereopod 1 (chelipeds): Dimorphic but spination than wide; regions indicated; dorsal surface armed with similar. Surfaces of both chelipeds smooth or with long, slender spines and scattered, widely separated scattered granules in addition to several small spines. granules. Gastric region convex, with 2 pairs of long, Coxa unarmed; with tufts of setae mesially. Ischiobasis upright spines. Cardiac region with pair of anterior with 2 or 3 blunt ventral spines and scattered tubercles. tubercles and posterior pair of long upright spines. Merus inner margin, with straight subdistal spine; Hepatic margin with low tubercle at midlength and ventral margin with scattered tubercles; dorsal mar- spine longest of carapace spines (0.58 pcl in holotype), gin with 5–7 small, scattered spines and 3 prominent directed anterolaterally. Intestinal region with pair of spines, increasing in length distally, distalmost shorter long upright spines on posterior margin. Branchial than half carpus length; lateral surface spinose, spines surface with 2 long upright spines and 2 conical tu- largest distally. Carpus with 6 spines and 3 or 4 small, bercles as follows: 1 spine at level of pereopod 3 coxa, scattered spines or acute tubercles. Palm with 13–15 1 spine at level of pereopod 4 coxa, 2 conical tubercles small scattered spines or tubercles on dorsal and lateral (or short spines in juveniles) in transverse row slightly surfaces, inner and ventral surfaces unarmed. behind posterior spine. Branchial margins spinose; Major cheliped 1.48–1.65 pcl (male), 1.39 (female); anterior branchial margin with 2 short spines, poste- upper palm length 1.17–1.42 times height (male), 1.42 rior spine the longer; 1st primary marginal branchial (female); occlusal margins of fingers corneous for distal spine about two-thirds length of hepatic spine; lateral fifth, proximally with 3 or 4 calcareous nodules; dacty- branchial margin with 1 or 2 short spines; 2nd primary lus dorsal margin broadly convex, with rows of tufts marginal branchial spine shorter than 1st primary mar- of golden setae and blunt proximal tooth, 1.22–1.24 ginal branchial spine; posterior branchial margin with times longer than dorsal margin of palm (male), 1.40 7 or 8 short spines. Pterygostomian region granulate or (female). tuberculate, with small, anterior, submarginal spine. Minor cheliped 1.54–1.60 pcl (male), 1.68 (female); Rostrum 0.71–0.82 pcl, comprising proximal and upper palm length 1.37–1.64 times height (male), 1.55 distal portions; proximal portion angled dorsally by (female); occlusal margin corneous for slightly less about 30°, with pair of long, divergent, dorsal spines than distal half, proximally crenulate; dactylus dorsal that overreach midlength of distal portion of rostrum; margin broadly convex, with rows of tufts of golden distal portion subhorizontal, distally bifurcate for setae and blunt proximal tooth, 1.54–1.60 times longer half length, forming pair of strongly divergent spines; than dorsal margin of palm (male), 1.68 (female). ventral rostral spine elongate, extending anteriorly Pereopods 2–4 (walking legs 1–3): Similar, slender, beyond apices of outer orbital spine. Posterior orbital spinose; surface between major spines with small, scat- margin concave; outer orbital spine directed anteri- tered spines. Pereopod 4 longest. Coxa of pereopods orly, slightly longer than anterolateral spine, reaching 2–4 unarmed. Ischiobasis dorsodistal margin with two slightly beyond cornea. small spines or blunt tubercles; ventrodistal margin Ocular peduncle: Longer than cornea; unarmed. with two spines, posteriormost spine of pereopod 4 Antennule: Peduncle unarmed, reaching anteriorly longest. Merus subcylindrical; extensor and flexor beyond antennal peduncle by full length of distal an- margins with small scattered spines and 3 or 4 major tennular peduncle article. spines in addition to long, prominent, distal spine, Antenna: Basal antennal article unarmed; outer mar- not exceeding half merus length; dorsal surface with gin of article 2 with long slender spine, not reaching irregular row of 6–10 spines of which 3 or 4 promi- midlength of article 4; article 3 unarmed; scaphocerite nent. Carpus surface with scattered, short spines and minute, blunt, shorter than article 4; article 4 unarmed, strongly elongated distal and second proximal spines, about half as long as article 5. each exceeding half carpus length. Propodus extensor Abdomen: Somite 1 with low triangular teeth on margins with 12–15 spines; flexor margins with 6–10

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 34. Lithodes rachelae sp. nov., female paratype, cl 80.0 mm, pcl 43.8 mm, cw 40.0 mm, SE of Albany, Western Australia (SAM C6853). A, dorsal habitus. C, ventral surface and abdomen.

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ spines. Dactylus curved, laterally compressed; proxi- the merus length:height ratio 7.6 versus a maximum mally with 2 spines on either side adjacent to articula- of 7.3. It should also be noted that the differences be- tion; extensor margin with 2–4 slender corneous spines tween L. rachelae and L. australiensis in relative length along length; flexor margin unarmed; apex corneous. and slenderness of the walking legs can be expected Pereopod 2 length 3.26–3.27 pcl (male), 2.85 pcl to be more pronounced when larger specimens of the (female). Merus 1.23–1.24 pcl (male), 1.01 pcl (female); former become available (largest known L. rachelae pcl length:height ratio 8.1–9.82 (male), 7.60 (female). 117.3 mm; largest L. australiensis pcl 133.8 mm), and Carpus 0.46–0.51 merus length (male), 0.49 (female). size-matched specimens can be directly compared. The Propodus 0.82–0.87 merus length (male), 0.85 (female); proportional length of the merus and overall length of length:height ratio 13.13–14.16 (male), 11.23 (female). the third walking leg of the smallest male L. rachelae (1.3 Dactylus 0.50–0.54 propodus length (male), 0.60 (fe- pcl and 3.6 pcl, respectively) exceeds that of the largest male). male L. australiensis (1.2 pcl; 3.5 pcl). In particular, more Pereopod 3 length 3.57–3.64 pcl (male), 3.06 pcl accurate comparisons between female L. australiensis (female). Merus 1.33–1.36 pcl (male), 1.07 pcl (female); and L. rachelae must await capture of adult females of length:height ratio 8.92–9.09 (male), 7.73 (female). both species. The female paratype of L. rachelae is a Carpus 0.47–0.51 merus length (male), 0.51 (female). subadult, with the abdomen not having reached full Propodus 0.85–0.87 merus length (male), 0.87 (female); size or maximum asymmetry. length:height ratio 14.72–16.61 (male), 11.84 (female). Lithodes rachelae is presently known only between Dactylus 0.48–0.52 propodus length (male), 0.57 (fe- the Great Australian Bight and Albany, Western Aus- male). tralia, whereas L. australiensis ranges from central New Pereopod 4 length 3.62–3.71 pcl (male), 3.10 pcl (fe- South Wales to Tasmania. male). Merus 1.29–1.30 pcl (male), 1.06 (female); length: One of two juvenile Lithodes, reported and figured height ratio 9.17–9.36 (male), 7.60 (female). Carpus as L. aff. longispina by Poore et al. (2008) was collected 0.51–0.55 merus length (male), 0.51 (female). Propodus from southeast of Albany, where adult L. rachelae is 0.88–0.91 merus length (male), 0.90 (female); length: known to occur. The juvenile is tentatively referred to height ratio 15.24–15.61 (male), 13.21 (female). Dactylus L. rachelae. The second specimen, from Perth Canyon 0.51–0.53 propodus length (male), 0.59 (female). (31°58.28’S, 115°06’W), was not available for examination at the time of the study. Colour in life. Deep-red overall (Pl. 1D). Distribution. South Australia, from the Great Aus- Etymology. Named for Rachel Ahyong, for her support tralian Bight, to southwestern Western Australia; during the course of the study. 930–1500 m.

Remarks. Lithodes rachelae sp. nov. most closely resem- Lithodes richeri Macpherson, 1990 bles L. australiensis sp. nov., L. robertsoni sp. nov., and (Figs 26, 35–37, Pl. 1E) L. paulayi Macpherson & Chan, 2008, sharing long carapace spines in adults; two long dorsal branchial Lithodes richeri. — Dawson, 1989: 317 [nomen nudum]. spines; a three-segmented abdominal somite 2 in Lithodes richeri Macpherson, 1990: 219–221, figs. 1a, 2b [type adults; and unarmed flexor margins of the dactyli of locality: New Caledonia]; 2001: 798–799. — Zaklan, 2002: the walking legs. Lithodes rachelae is readily separated 767, 782. — ?Takeda & Nagai, 2004: 15–18, figs 5, 6. from L. paulayi in having proportionally longer primary branchial spines, with the 2nd primary branchial spine Type material. Holotype: MNHN Pg 4269, male (cl 166.5 in the new species distinctly longer, rather than shorter mm, pcl 89.0 mm, cw 81.0 mm), New Caledonia, out- than the outer oribital spine; and in longer walking legs side coral reef, trap. with the third leg of males 3.6–3.7 versus less than 3.2 pcl. Lithodes rachelae differs from L. australiensis and Other material examined. Loyalty Islands: IRD, 2 males L. robertsoni in having proportionally longer, more (cl 179.5 mm, pcl 95.3 mm, cw 87.4 mm; pcl 106.6 mm, slender walking legs, being most pronounced in adult cw 100.0 mm), off Lifou, 20°54’N, 166°59’E, 980–1000 males in which the merus of the third walking leg is 1.3 m, 17–18 Jun 1977. pcl versus less than 1.2 pcl in the latter two species (fur- Australia: AM P40778, 1 male (cl 100.9 mm, pcl 53.6 ther features distinguishing L. rachelae from L. robertsoni mm, cw 49.6 mm), E of Tuncurry, 32°02’S, 153°09’E, are given under the account of the latter). The third 1025–1080 m, FRV Kapala, K88-08-04, coll. K. Graham, walking leg of male L. rachelae is 3.6–3.7 versus less than 4 May 1988; AM P53233, 1 male (pcl 42.2 mm, cw 36.8 3.5 pcl, with a merus length:height ratio 9.2–9.4 versus mm), E of Cape Hawke, 32°06’S, 153°08’E, 952–970 m, a maximum of 7.8 in L. australiensis. The third walking FRV Kapala, K84-10-06, coll. K. Graham, 18 Jul 1984; AM leg of female L. rachelae is 3.1 versus 2.6 pcl or less and P35600, 1 male (cl 205.0 mm, pcl 112.1 mm, cw 105.8

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 35. Lithodes richeri Macpherson, 1990, male, pcl 53.6 mm, E of Tuncurry (AM P40778). A, dorsal habitus. B, ventral surface and abdomen. C, posterior carapace and abdominal somite 2.

mm), E of Cape Hawke, 32°06’S, 153°08’E, 952–970 m, nile females (pcl 27.4–52.6 mm, cw 24.2–46.3 mm), E FRV Kapala, K84-10-06, coll. K. Graham, 18 Jul 1984; AM of Tuncurry, 32°08’S, 153°09’E, 1080 m, FRV Kapala, P35593, 1 male (cl 99.0 mm, pcl 50.5 mm, cw 45.4 mm), K89-12-04, coll. K. Graham, 15 Jun 1989; AM P75312, 1 E of Cape Hawke, 32°08–04’S, 153°09–10’E, 1006–1052 ovigerous female (cl 255.0 mm, pcl 143.0 mm, cw 127.0 m, FRV Kapala, K84-10-04, coll. K. Graham; AM P52704, mm), E of Broken Bay, 33°35’S, 152°05’E, 860 m, from 4 males (pcl 57.6–97.3 mm, cw 52.7–90.1 mm), 3 juve- fisherman, 6 Jan 1992; AM P35594, 1 male (cl 56.2 mm,

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ pcl 26.8 mm, cw 23.7 mm), E of Shoalhaven Heads, spine short, little longer than anterior branchial mar- 34°51–55’S, 151°15–14’E, 1042–1061 m, FRV Kapala, ginal spines; posterior branchial margin with 5–7 short K83-14-05, coll. K. Graham, 26 Oct 1983; AM P35597, spines. All marginal branchial spines reduced to small 4 males (pcl 21.6–25.2 mm, cw 18.8–23.1 mm), E of protrusions in largest specimens. Pterygostomian Shoalhaven Heads, 34°53–56’S, 151°15–13’E, 1079–1116 region granulate or tuberculate, with small, anterior, m, FRV Kapala, K83-14-06, coll. K. Graham, 26 Oct 1983; submarginal spine. AM P35602, 1 female (cl 210.2 mm, pcl 101.3 mm, cw Rostrum as slightly shorter to slightly longer than 94.4 mm), E of Brush Island, 35°28–31’S, 150°54–50’E, pcl (0.82–1.08 pcl), comprising proximal and distal 905–960 m, FRV Kapala, K84-11-01, coll. K. Graham, portions; proximal portion angled dorsally by at least 26 Jul 1984; AM P35605, 1 ovigerous female (cl 131.4 30°; with pair of divergent, dorsal spines at rostral mi- mm, pcl 125.1 mm, cw 132.9 mm), E of Brush Island, dlength; distal portion subhorizontal, distally bifurcate 35°38’S, 150°44’E, 887 m, FRV Kapala, K83-11-03, coll. for half to two-thirds length, forming pair of strongly K. Graham, 8 Sep 1983. divergent spines; ventral rostral spine elongate, extending anteriorly beyond apices of outer orbital spine. Diagnosis. Carapace dorsal surface and margins with Posterior orbital margin concave; outer orbital spine slender spines, becoming reduced to short conical pro- directed anteriorly, subequal to or shorter than ante- tuberances in large adults; gastric region with 4 upright rolateral spine, not reaching beyond cornea. spines, anterior pair longer than half length of hepatic Ocular peduncle: Longer than cornea; unarmed. spine, posterior pair half length of hepatic spine; car- Antennule: Peduncle unarmed, reaching anteriorly diac and intestinal regions each with 2 upright spines; beyond antennal peduncle by three-quarters to full branchial surface with 2 long upright spines. First pri- length of distal antennular peduncle article. mary marginal branchial spine half length of hepatic Antenna: Basal antennal article unarmed; outer mar- spine or less, 2nd primary marginal branchial spine gin of article 2 with slender spine, not reaching beyond minute, not larger than posterior branchial spines; midlength of article 4; article 3 unarmed; scaphocerite posterior branchial margin with 5–7 short spines or minute, blunt, shorter than article 4; article 4 unarmed, protuberances. Rostrum exceeding 0.8 pcl; proximal about half as long as article 5. half strongly upraised by about 30°. Abdominal somite Abdomen: Somite 1 with low triangular teeth on pos- 2 comprising single fused plate. Antennal peduncle terior margin. Somite 2 rugose; plates fused into single article 2 with slender outer spine not reaching beyond unit; median surface with pair of submedian spines or midlength of article 4. Chelipeds unequal; palms with angular prominences; posterior margin with 4–6 stout low tubercles or spines on dorsal surface. Walking legs spines or blunt angular prominences; lateral margins spinose; surface between major spines smooth or with irregularly dentate. Somites 3–5 with nodular median very few, scattered spines. Pereopod 4 merus longer plates; surface of submedian plates irregular, unarmed; than pcl in both sexes; dactylus flexor margin unarmed, marginal plates of males subdivided, with angular half propodus length. or sharp apices; marginal plates of females with left margin smooth to crenulate (fused with submedians), Description. Carapace: Pyriform, 1.07–1.15 times longer right marginals subdivided, with rounded or pointed than wide; regions indicated; dorsal spines slender, apices. Somite 6 of both sexes longer than wide, distal becoming reduced to short conical protrusions in large margin with 2 or 4 distal spines or prominences. Telson adults; dorsal surface otherwise smooth apart from rounded, unarmed. scattered, widely separated granules. Gastric region Pereopod 1 (chelipeds): Dimorphic but spination convex, with 2 pairs of spines. Cardiac region with similar. Surfaces of both chelipeds smooth or with pair of anterior tubercles and posterior pair of spines. scattered granules in addition to major spines. Coxa Hepatic spine longest of carapace spines (0.1 pcl in with tufts of setae, unarmed; ischiobasis with 3 stout 89.0 mm male holotype), directed anterolaterally. In- ventral spines and about 6 blunt tubercles. Merus inner testinal region with pair of spines on posterior margin. margin, with stout subdistal spine and 2 or 3 smaller Branchial surface with 2 spines and 2 low tubercles as spines or acute tubercles; ventral margin with two rows follows: 1 spine at level of pereopod 3 coxa, 1 spine at of 2 or 3 low conical spines; dorsal and lateral surface level of pereopod 4 coxa, 2 conical tubercles in trans- sparsely spinose, spines largest distally. Carpus with verse row slightly behind posterior spine. Branchial 2 or 3 short spines on dorsal margin; lateral surfaces margins spinose; anterior branchial margin with 2 with distal spine and scattered acute tubercles; ventral short spines, posterior spine subeqal to or longer than margin with small, scattered, acute tubercles. Palms of anterior; 1st primary marginal branchial spine half as both chelipeds in both sexes with similar ornamenta- long as hepatic spine or less; lateral branchial margin tion; with small spines and acute tubercles on dorsal, with 2 or 3 short spines; 2nd primary marginal branchial lateral, and ventral surfaces, inner surface with few

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 36. Lithodes richeri Macpherson, 1990. A–C, female, pcl 101.3 mm, E of Brush Island (AM P35602). D, male, pcl 53.6 mm, E of Tuncurry (AM P40778). A, dorsal habitus. B, D, carapace, right lateral view. C, ventral surface and abdomen.

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ low tubercles; dorsal margin with row of 3 or 4 small (male), 13.12–13.17 (female). Dactylus 0.45–0.52 pro- conical spines; midlateral surface and ventral surfaces podus length (male), 0.46–0.49 (female). with small scattered tubercles or small spines. Pereopod 4 length 3.54–3.81 pcl (male), 3.04–3.47 Major cheliped 1.25–1.38 pcl (male), 1.21–1.32 pcl (female). Merus 1.31–1.38 pcl (male), 1.10–1.25 (female); upper palm length 1.25–1.48 times height pcl (female); length:height ratio 9.28–9.96 (male), (male), 1.24–1.27 (female); occlusal margins of fingers 8.35–8.42 (female). Carpus 0.44–0.51 merus length corneous for distal third, proximally with 3 calcareous (male), 0.50–0.51 (female). Propodus 0.81–0.91 merus nodules; dactylus dorsal margin broadly convex, with length (male), 0.87–0.88 (female); length:height ratio rows of tufts of golden setae and blunt proximal tooth, 14.60–15.46 (male), 12.60–12.72 (female). Dactylus 0.50 1.31–1.79 times longer than dorsal margin of palm propodus length (male), 0.46–0.49 (female). (male), 1.46–1.47 (female). Minor cheliped 1.21–1.35 pcl (male), 1.22–1.28 Colour in life. Overall deep-red (Pl. 1E). (female); upper palm length 1.41–1.60 times height Remarks. Poore’s (2004) suggestion that previous re- (male), 1.33–1.41 (female); occlusal margin corneous ports of L. longispina from eastern Australia (Dawson & for slightly less than distal half, proximally crenulate; Yaldwyn 1985) could be based on L. richeri is confirmed dactylus dorsal margin broadly convex, with rows of by the present series. tufts of golden setae and blunt proximal tooth, 1.75– The Australian specimens of L. richeri agree well 1.96 times longer than dorsal margin of palm (male), with New Caledonian material and exhibit morpho- 1.88–1.90 (female). logical variation as reported by Macpherson (1990). As Pereopods 2–4 (walking legs 1–3): Similar, slender, in the holotype of L. richeri, the marginal and dorsal elongate; ornamentation similar in both sexes; seg- carapace spines of the largest Australian specimens are ments sparsely spinose, surface between major spines reduced or obsolete, but longer in smaller individuals. smooth or with few, scattered spines or granules. A distinctive feature of L. richeri is the considerably Pereopod 4 longest. Distal margins of coxae unarmed; elongated rostrum, measuring almost as long as or surface smooth. Ischiobasis with 3 or 4 non-setose longer than the pcl; the rostrum in adults of other Aus- distal spines. Merus subcircular to ovate in cross sec- tralian and New Zealand lithodids is always distinctly tion; extensor margin with 4–10 spines of which 2 or 3 shorter than pcl. The dorsal spines in L. richeri are long distinctly longer in addition to prominent distal spine; and slender in juveniles, becoming reduced to short, flexor margin with 2 rows of 4 or 5 small spines; up- conical protrusions in large adults. The hepatic spine per surface with few, small, scattered spines. Carpus is as long as the pcl in the 26.8 mm male (AM P35594), about half merus length; extensor margin with distal about half pcl in the 53.6 mm male (AM P40778), and and second proximal spines longest, not exceeding about 0.1 pcl in the 89.0 mm male holotype. In the 143 one-third carpus length in adults; surface with few, mm female (AM P75312), the hepatic spine is less than small, scattered spines. Propodus ovate in cross sec- 0.05 pcl. tion, with 6–9 small spines on extensor margin and Lithodes richeri is sometimes sympatric with Lithodes few small spines on dorsal surface; flexor margin with australiensis and distinguishing between juveniles of 9–15 small spines. Dactylus curved, rounded in cross the two species can be difficult. Adults are readily section distally; proximally with 2 spines on either side separated by the marked difference in rostral length adjacent to articulation; with 0–4 small corneous spines (almost as long as pcl in L. richeri; less than half pcl in along extensor margin; flexor margin smooth or with L. australiensis), marked difference in dorsal spine few minute, widely spaced setae; apex corneous. length (reduced to short conical projections in Pereopod 2 length 2.96–3.17 pcl (male), 2.47–2.91 L. richeri; prominent in L. australiensis), slenderness pcl (female). Merus 1.13–1.16 pcl (male), 0.91–1.07 pcl of the walking legs (pereopod 4 merus length:height (female); length:height ratio 8.73–9.54 (male), 7.35–7.58 ratio of males exceeds 9 in L. richeri; less than 8 in (female). Carpus 0.44–0.50 merus length (male), 0.50– L. australiensis), spination of the branchial margins of 0.52 (female). Propodus 0.80–0.87 merus length (male), the carapace (at most one prominent primarly marginal 0.84–0.85 (female); length:height ratio 13.24–13.88 branchial spine in L. richeri; two in L. australiensis), and (male), 11.44–12.34 (female). Dactylus 0.49–0.55 pro- fusion of the plates of abdominal somite 2 (fused in podus length (male), 0.50–0.51 (female). L. richeri; separate in L. australiensis). Juvenile L. richeri Pereopod 3 length 3.36–3.57 pcl (male), 2.82–3.29 and L. australiensis are more difficult to separate be- pcl (female). Merus 1.29–1.31 pcl (male), 1.03–1.22 pcl cause of their similarly long dorsal spines and rostra. (female); length:height ratio 9.14–9.79 (male), 8.08–8.17 As in adults, however, the fused plates of abdominal (female). Carpus 0.43–0.48 merus length (male), 0.48– somite 2 and the presence of only one prominent pri- 0.50 (female). Propodus 0.78–0.86 merus length (male), mary marginal branchial carapace spine will separate 0.85–0.87 (female); length:height ratio 13.99–14.87 juvenile L. richeri from L. australiensis. In addition, the

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 37. Lithodes richeri Macpherson, 1990. Carapace. A, juvenile male, pcl 26.8 mm, E of Shoalhaven Heads (AM P35594). B, female, pcl 143.0 mm, E of Broken Bay (AM P75312). C, male, pcl 53.6 mm, E of Tuncurry (AM P40778). D, female, pcl 101.3 mm, E of Brush Island (AM P35602).

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ flexor margins of the walking leg dactyli of juvenile mm, cw 113.3 mm), 39°45.97–46.29’S, 167°15.49–15.14’E, L. australiensis up to about 60 mm pcl are armed with a 880–1044 m, FV Thomas Harrison, THH0601/15, 26 Jun row of corneous spinules, but are unarmed in L. richeri 2006. of all sizes examined (smallest with pcl 21.6 mm). Takeda & Nagai (2004) tentatively identified a Diagnosis. Carapace dorsal surface with slender spines large female specimen (pcl 175 mm) from the Ceram in adults; gastric region with 4 upright spines, anterior Sea, Indonesia, as L. richeri, which differed from the pair longer than half length of hepatic spine, posterior type description in carapace shape — the carapace pair half length of hepatic spine; cardiac and intestinal of the Ceram specimen is widest across the anterior, regions each with 2 long upright spines; branchial surface with 2 long upright spines. Branchial margins rather than mid-portion of the branchial region, with st angular rather than evenly rounded anterior branchial spinose; 1 primary marginal branchial spine about half length of hepatic spine, 2nd primary marginal branchial margins, and the junction of the hepatic and branchial st regions forming an angular rather than evenly curved spine shorter than 1 primary marginal branchial spine; concavity. Takeda & Nagai (2004) speculated that the posterior branchial margin with 7 or 8 short spines and differences in carapace shape might be attributable 1 or 2 low tubercles. Rostrum not exceeding 0.8 pcl, to allometry, with the Ceram specimen considerably upraised by about 30°; almost straight or only slightly larger than the holotype (pcl 81 mm). The specimens angled near midlength. Abdominal somite 2 compris- in series of L. richeri reported here, however, span a ing 3 plates in adults. Antennal peduncle article 2 wide size range (pcl 26.8–143.0 mm), and the carapace with slender outer spine not reaching beyond article outline is consistently pyriform, exhibiting no trend 4. Chelipeds unequal; palms with low tubercles or towards the condition of the Ceram specimen. Addi- spines on dorsal surface. Walking legs spinose; surface tionally, the dorsal carapace spines are distinctly more between major spines with very few, scattered spines. prominent than in the largest specimen examined here Pereopod 4 merus 1.1 pcl or less, length less than 8.2 (pcl 143.0 mm). The Ceram specimen possibly repre- times height; propodus length 11–14 times height sents an undescribed species. (male), 11 (female); dactylus flexor margin unarmed. Although Australian records of L. richeri are limited Description. Carapace: Pyriform, 1.03–1.09 times longer to New South Wales, it can be expected to occur off than wide; regions indicated; dorsal surface armed Queensland given its occurrence at Vanuatu, Lifou, with long, slender spines and numerous, well-spaced and New Caledonia. granules. Gastric region convex, with 2 pairs of long, upright spines. Cardiac region with pair of anterior Distribution. Vanuatu, Lifou, New Caledonia, and eastern Australia south to at least 35°S; possibly from tubercles and posterior pair of long upright spines. the Ceram Sea, Indonesia; 860–1220 m (Macpherson, Hepatic spine longest of carapace spines (0.3 pcl in 2001; present study). holotype; 0.5 pcl in 70.2 mm pcl female paratype), directed anterolaterally. Intestinal region with pair of long upright spines on posterior margin. Branchial Lithodes robertsoni sp. nov. surface with 2 long upright spines and 2 small conical (Figs 31, 38–42, Pl. 1F) spines as follows: 1 spine at level of pereopod 3 coxa, 1 Type material. Holotype: NIWA 42889, male (cl 186.6 spine at level of pereopod 4 coxa, 2 small conical spines mm, pcl 128.1 mm, cw 117.0 mm), near Pitt Island, in transverse row slightly behind posterior spine. Chatham Rise, 44°41.6’S, 177°01.4’W, 1002 m, trip Branchial margins spinose; anterior branchial margin 2521/69, FV Thomas Harrison, coll. T. Turton, 15 Nov with 2 small spines, subequal or posterior spine longer 2007. than anterior; 1st primary marginal branchial spine Paratypes: NIWA 34914, 1 male (cl 122.7 mm, pcl 81.7 about half length of hepatic spine; lateral branchial mm, cw 76.4 mm), The Snares, 48°02.01’S, 166°06.00’E, margin with 1 short spine and 1 short granule or 3 935 m, trip 1171/12, Z9583, 25 Nov 1998; NIWA 34908, spines (NIWA 60564); 2nd primary marginal branchial 1 female (cl 119.0 mm, pcl 71.2 mm, cw 65.2 mm), spine shorter than two-thirds length of 1st primary mar- Bounty Trough, 46°27.0’S, 172°01.8’E, 1030–1236 m, trip ginal branchial spine; posterior branchial margin with 1650, tow 58, FV Amital Explorer, 18 May 2002; NIWA 7 or 8 spines and 1 or 2 low tubercles. Pterygostomian 61239, 1 female (pcl 70.2 mm, cw 64.6 mm), Bounty region granulate or tuberculate, with small, anterior, Trough, 47°53.1’S, 175°02.5–02.7’E, 1252–1259 m, trip submarginal spine. 2494, tow 27, 4 Sep 2007. Rostrum half to two-thirds pcl, upraised by about 30°; almost straight or only slightly angled near mi- Other material examined. Chatham Rise: NIWA 60564, dlength; with pair of long, divergent, dorsal spines 1 badly damaged male (pcl 103.7 mm, cw 100.2 mm), that overreach midlength of distal portion of rostrum; Chatham Rise, FV San Waitaki, SWA0901/-. Challenger distally bifurcate for about half length, forming pair Plateau: NIWA 62808, 1 badly damaged male (pcl 114.2 of strongly divergent spines; ventral rostral spine

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 38. Lithodes robertsoni sp. nov., male holotype, cl 186.6 mm, pcl 128.1 mm, cw 117.0 mm, Chatham Rise (NIWA 42889). A, dorsal habitus. B, carapace, right lateral view.

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 39. Lithodes robertsoni sp. nov., male holotype, cl 186.6 mm, pcl 128.1 mm, cw 117.0 mm, Chatham Rise (NIWA 42889). A, carapace, dorsal view. B, anterior carapace, dorsal view. C, abdomen. D, posterior carapace and abdominal somite 2.

elongate, extending anteriorly beyond apices of outer Antenna: Basal antennal article unarmed; outer mar- orbital spine. Posterior orbital margin concave; outer gin of article 2 with slender spine, not reaching apex of orbital spine directed anteriorly, slightly longer than article 4; article 3 unarmed; scaphocerite minute, blunt, anterolateral spine, reaching to or beyond cornea. shorter than article 4; article 4 unarmed, about half as Ocular peduncle: Longer than cornea; unarmed. long as article 5. Antennule: Peduncle unarmed, reaching anteriorly Abdomen: Somite 1 with low triangular teeth on beyond antennal peduncle by about three-quarters posterior margin. Somite 2 rugose; composed of 3 length of distal antennular peduncle article. distinct plates separated by narrow groove; median

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 40. Lithodes robertsoni sp. nov., male holotype, cl 186.6 mm, pcl 128.1 mm, cw 117.0 mm, Chatham Rise (NIWA 42889). A, right chela. B, left chela. C, left pereopod 4. D, left antenna and orbit. E, anterior carapace, right lateral view. F, carapace, left posterior.

surface with pair of stout submedian spines; posterior irregular, unarmed except for 1 or 2 small distolateral margin with 4–8 stout teeth or angular prominences; teeth; marginal plates of males subdivided, with an- lateral margins irregularly dentate. Somites 3–5 with gular or sharp apices; marginal plates of females with nodular median plates; surface of submedian plates left margin spinose (fused with submedians), right

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ marginals subdivided, with angular to spiniform small spines on dorsal surface; flexor margin with 4–7 apices. Somite 6 of both sexes longer than wide, distal small spines. Dactylus curved, rounded in cross sec- margin with 4 distal spines or prominences. Telson tion distally; proximally with 2 spines on either side rounded, unarmed. adjacent to articulation; with 3 or 4 small corneous Pereopod 1 (chelipeds): Dimorphic but spination spines along extensor margin; flexor margin smooth similar. Surfaces of both chelipeds smooth or with or with about 4 soft, minute, widely spaced setae; apex scattered granules in addition to major spines. Coxa corneous. with tufts of setae, unarmed; ischiobasis with 3 stout Pereopod 2 length 2.76–2.95 pcl (male), 2.63–2.66 ventral spines and about 6 blunt tubercles. Merus inner pcl (female). Merus 0.99–1.07 pcl (male), 0.90–0.92 pcl margin with stout subdistal spine and 2 or 3 smaller (female); length:height ratio 6.26–7.50 (male), 5.70–6.15 spines or acute tubercles; ventral margin with two (female). Carpus 0.51–0.53 merus length (male), rows of 2 or 3 low conical spines; dorsal and lateral 0.52–0.54 (female). Propodus 0.84–0.85 merus length surface spinose, spines largest distally. Carpus with (male), 0.86–0.90 (female); length:height ratio 10.26– prominent spines on dorsal and lateral surfaces; dorsal 13.22 (male), 9.89–10.71 (female). Dactylus 0.52–0.54 margin with row of 3 stout spines; ventral margin with propodus length (male), 0.59–0.62 (female). small, scattered, acute tubercles. Palms of both cheli- Pereopod 3 length 3.04–3.18 pcl (male), 2.84–2.87 peds in both sexes with similar ornamentation; with pcl (female). Merus 1.08–1.13 pcl (male), 0.97–0.98 pcl small spines and acute tubercles on dorsal, lateral and (female); length:height ratio 6.92–7.87 (male), 6.05–6.58 ventral surfaces, inner surface with few low tubercles; (female). Carpus 0.52 merus length (male), 0.53–0.55 dorsal margin with row of 3 or 4 small conical spines; (female). Propodus 0.85–0.86 merus length (male), midlateral surface with 2 rows of 3 or 4 small spines 0.89–0.90 (female); length:height ratio 10.78–13.76 or tubercles, of similar size to dorsal row; ventral sur- (male), 10.71–11.37 (female). Dactylus 0.48–0.52 pro- face with scattered tubercles, smaller than lateral and podus length (male), 0.57–0.61 (female). dorsal spines. Pereopod 4 length 3.00–3.21 pcl (male), 2.91–2.92 Major cheliped 1.27–1.52 pcl (male), 1.33–1.34 pcl (female). Merus 1.05–1.10 pcl (male), 0.96–0.98 pcl (female); upper palm length 1.22–1.35 times height (female); length:height ratio 6.70–8.16 (male), 6.11–6.76 (male), 1.29–1.40 (female); occlusal margins of fingers (female). Carpus 0.55 merus length (male), 0.55–0.58 corneous for distal quarter, proximally with 3 calcare- (female). Propodus 0.90–0.92 merus length (male), ous nodules; dactylus dorsal margin broadly convex, 0.92–0.95 (female); length:height ratio 11.43–14.08 with rows of tufts of golden setae and blunt proximal (male), 11.55–11.87 (female). Dactylus 0.51–0.53 pro- tooth, 1.18–1.39 times longer than dorsal margin of podus length (male), 0.54–0.62 (female). palm (male), 1.32–1.36 (female). Minor cheliped 1.36–1.50 pcl (male), 1.32–1.33 Colour in life. Overall deep-red (Pl. 1F). (female); upper palm length 1.36–1.53 times height (male), 1.43–1.52 (female); occlusal margin corneous Etymology. Named for Don Robertson, NIWA, on oc- for slightly less than distal half, proximally crenulate; casion of his retirement, and for his strong support and dactylus dorsal margin broadly convex, with rows of advocacy for biodiversity research. tufts of golden setae and blunt proximal tooth, 1.61– 1.70 times longer than dorsal margin of palm (male), Remarks. Lithodes robertsoni sp. nov. closely resembles 1.68–1.73 (female). L. australiensis sp. nov., L. rachelae sp. nov., and Pereopods 2–4 (walking legs 1–3): Similar, slender, L. paulayi Macpherson & Chan, 2008, species sharing elongate; ornamentation similar in both sexes; seg- long dorsal spines in adults, two long dorsal branchial ments spinose, surface between major spines with spines, unarmed flexor margins of the walking leg minute granules and few, scattered spines. Pereopod 3 dactyli, and a three-segmented abdominal somite 2 and 4 subequal, longer than pereopod 2. Distal margins in adults. Unfortunately, the aforementioned species of coxae unarmed; surface smooth. Ischiobasis with 3 or exhibit few meristic differences, and are best separated 4 non-setose distal spines. Merus subcircular to ovate via morphometrics of the walking legs. The walking in cross section; extensor margin with 7–10 spines of legs of L. robertsoni are proportionally shorter and less which 2 or 3 are distinctly longer in addition to long, slender than in L. rachelae. The overall pereopod 4 prominent, distal spine; flexor margin with 2 rows of length in L. robertsoni does not exceed 3.2 pcl, compared 5–7 small spines; upper surface with irregular row of to 3.6 in male L. rachelae; and in females, less than 3.0 7–9 small spines. Carpus half merus length of merus; versus 3.1 or greater, respectively. The pereopod 4 extensor margin with distal and second proximal merus length of male L. robertsoni is shorter than 1.1 spines longest, exceeding half carpus length; surface pcl with a length:height ratio <8.2 versus about 1.3 with scattered small spines. Propodus compressed; pcl and >9.1, respectively, in male L. rachelae. Lithodes with 7–11 small spines on extensor margin and 4–7 robertsoni can be readily distinguished from L. paulayi

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 41. Lithodes robertsoni sp. nov., female paratype, pcl 70.2 mm, cw 64.6 mm, Bounty Trough (NIWA 61239). A, dorsal habitus. B, abdomen.

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 42. Lithodes robertsoni sp. nov. A, male, pcl 103.7 mm, Chatham Rise (NIWA 60564). B, male paratype, cl 122.7 mm, pcl 81.7 mm, cw 76.4 mm, The Snares (NIWA 34914).

by the longer outer spine on antennal article 2, which for 128.1 mm and 133.8 mm specimens of L. robertsoni reaches to, or almost to, the end of article 4, rather and L. australiensis, respectively). than to the midlength in L. paulayi; and the less slender Mature female L. robertsoni and L. australiensis are pereopod 4 propodi (length 11–14 times height versus currently known only from very few specimens, and no about 17). large females of either species have yet been collected. Lithodes robertsoni is morphologically closest to Based on current data, the propodus of pereopod 4 of L. australiensis. The two species are difficult to dis- female L. robertsoni appears to be slightly more slender tinguish and it is with some hesitation that both are than in L. australiensis (length exceeding 11.5 times recognised as separate new species. Comparisons height versus about 11). Unfortunately, L. robertsoni is are confounded by the limited available material of apparently an uncommon species, so its full range of L. robertsoni, in which the largest and most complete morphological variation remains to be documented. specimen is the male holotype. The two forms, however, Moreover, most specimens are damaged and have exhibit subtle but consistent morphological differences broken rostra. More ‘satisfactory’ characterisation of in addition to disjunct distributions (southeastern New differences between L. robertsoni and L. australiensis Zealand versus southeastern Australia). The number of must await capture of more specimens of both species, posterior marginal branchial spines occupies a higher especially females. A badly damaged male (NIWA range in L. robertsoni (7 or 8) than in L. australiensis (5 60564, pcl 103.7 mm), with a broken rostrum and or 6), but occasional overlap seems likely given the broken carapace spines, and with only the proximal closeness of the ranges. Adult, size-matched male segments of the walking legs intact, is tentatively as- L. robertsoni differ from adult male L. australiensis in signed to L. robertsoni. It agrees well with other males having proportionally shorter walking legs. Pereopod in most aspects that can be studied, but differs in hav- 4 is 3.0 pcl at 81.7 mm pcl in L. robertsoni compared to ing three stout spines, instead of one, on the lateral 3.1 pcl in 78.0 mm pcl L. australiensis. At 128.1 mm pcl, branchial margin. pereopod 4 is 3.2 times pcl in L. robertsoni compared to In New Zealand waters, the only species that L. 3.5 for L. australiensis at 133.8 mm pcl. The pereopod 4 robertsoni is likely to be mistaken for is L. aotearoa, merus is proportionally shorter in L. robertsoni than in especially because the geographic and bathymetric L. australiensis (1.05–1.10 pcl vs 1.10–1.17 pcl across the ranges of the species overlap. Adult L. robertsoni dif- adult size range), with the largest specimens exbiting fers from L. aotearoa in abdominal somite 2 compris- the most marked differences (1.10 pcl versus 1.17 pcl ing three separate plates rather than a single fused

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ plate; in usually having only one spine and one or N. diomedeae (Benedict, 1895) [southeastern Pacific to two granules on the lateral branchial margin rather southwestern Atlantic oceans] than two or three spines; the surface of the submedian N. duhameli Macpherson, 2004 [Crozet Islands, south- plates on the male abdominal somites 2–5 do not bear western Indian Ocean] acute tubercles except distolaterally; the pereopod 4 N. flindersi Ahyong, 2010 [southeastern Australia, merus is always longer than the pcl in males rather southwestern Pacific Ocean and Southern Ocean] than usually shorter than pcl in L. aotearoa (except in N. grimaldii (A. Milne-Edwards & Bouvier, 1894a) some of the largest specimens); and the dorsal spines [northern Atlantic Ocean] are proportionally longer. By 120 mm pcl, the dorsal N. nipponensis Sakai, 1971 [Japan to Taiwan, northwest- spines are reduced to short conical protrusions in ern Pacific Ocean] L. aotearoa of about 0.1 pcl, rather than slender spines of N. vinogradovi Macpherson, 1988b [southeastern Indian at least 0.2 pcl in L. robertsoni. Lithodes robertsoni might Ocean] also mature at a smaller size than L. aotearoa. The 71.2 N. yaldwyni Ahyong & Dawson, 2006 [Ross Sea, Ant- mm pcl female of L. robertsoni (NIWA 34908) appears arctica] to be mature, based on abdomen development. Lithodes aotearoa matures by about 90 mm pcl. Remarks. Neolithodes currently includes 12 species including N. bronwynae sp. nov., described herein Distribution. New Zealand, from the Challenger Pla- from New Zealand. Five species are known from teau, Chatham Rise and Bounty Trough to The Snares; the Atlantic Ocean, of which two also range into the 935–1259 m. southeastern Pacific and southwestern Indian Ocean, respectively. Two species (N. vinogradovi Macpherson, Neolithodes A. Milne-Edwards & Bouvier, 1894 1988b, N. duhameli Macpherson, 2004) are known from the southern Indian Ocean, two from New Zealand Neolithodes A. Milne-Edwards & Bouvier, 1894b: 120 [type (N. brodiei Dawson & Yaldwyn, 1970 and N. bronwynae species: Lithodes grimaldii A. Milne-Edwards & Bouvier, sp. nov.), one from the Ross Sea Antarctica (N. yald- 1894a, by monotypy. Gender: masculine]. wyni Ahyong & Dawson, 2006), one from southern Australia (N. flindersi Ahyong, 2010), and one from Diagnosis. Carapace pyriform, not covering bases of walking legs; regions indicated; gastric region elevated the northwestern Pacific (N. nipponensis Sakai, 1971). above other regions; cardiac region triangular, separat- Records assigned to known Neolithodes species from ed from gastric region by deep groove; cervical groove other localities, especially in the Indo-Pacific, require shallow, indistinct. Rostrum comprised of median verification. In particular, records ofN . agassizi (Smith, spine and one pair of dorsal spines at base. Abdominal 1882) from the northwestern Indian Ocean (Anderson somite 2 comprising 5 plates (median plate, paired 1896; Alcock & Anderson 1899; Alcock 1901) are based submedian and paired marginal plates). Abdominal on an undescribed species (Dawson 1989). Similarly, somites 3–5 of males composed of small spiniform records of N. nipponensis from the southwestern Pacific nodules embedded in arthrodial membrane; female and records of N. brodiei from Vanuatu appear to be somites composed either of well-developed plates or based on undescribed species. Records of N. vinogradovi spiniform nodules on right side, and well-developed from New Caledonia appear referable to N. bronwynae. plates on left side. Sternite 5 (between pereopods 2) Macpherson (2001) recorded an unidentifiable juvenile with deep median fissure. Scaphocerite absent or a Neolithodes from the Marquesas Islands (French Poly- simple spine. Walking legs (pereopods 2–4) similar in nesia); as yet, adult Neolithodes have not been identified form, third walking leg longest, always longer than from the central Pacific. pcl; dactyli of adults without row of corneous flexor Neolithodes includes some of the deepest occurring spines. lithodids, down to 3200 m for N. capensis (see Macpher- son 1988b). Composition Key to species of Neolithodes from New Zealand, N. agassizi (Smith, 1882) [western Atlantic Ocean] Australia, and the Ross Sea N. asperrimus Barnard, 1947 [southeastern Atlantic Ocean to southwestern Indian Ocean] 1. Carapace surface between major spines with N. brodiei Dawson & Yaldwyn, 1970 [New Zealand, sparsely distributed granules or secondary spinules. southwestern Pacific Ocean] Pereopod 2–4 merus subcylindrical...... 2 N. bronwynae sp. nov. [New Zealand and Tasman Sea, southwestern Pacific Ocean] – Carapace surface between major spines densely N. capensis Stebbing, 1905 [South Africa, western In- covered with granules or secondary spinules. Pere- dian Ocean] opod 2–4 merus compressed, flattened...... 3

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ 2. Carapace covered with long, slender dorsal spines 78.4 mm), 38°33.6’S, 167°00.2’E, 1168–1179 m, FV Ar- in adults; longest spines exceeding 0.2 pcl [New row A02/143/83, 16 Sep 1983; NMNZ Cr7351, 1 male Zealand, northern Tasman Sea]...... (pcl 124.7 mm, cw 121.5 mm), W Challenger Plateau, ...... N. bronwynae sp. nov. 39°51.90’S, 167°09.00’E, 996 m, FV Oyang 7, trip 284, coll. Sharples, 9 Sep 1988; NMNZ Cr3746, 1 male (pcl – Carapace spines short, conical in adults; longest 137.5 mm, cw 136.6 mm), 39°27.9’S, 167°13.6’E, 1153– spines shorter than 0.2 pcl [Ross Sea, Antarctica].. 1155 m, FV Arrow, A02/108/83, 16 Sep 1983; NIWA ...... N. yaldwyni 31729, 1 juvenile, 39°55.57’S, 167°41.60’E, 1139–1144 m, 3. Pereopod 2–4 coxae of males and juvenile females TAN0707/67, 1 Jun 2007; NMNZ Cr3748, 1 male (pcl covered with short conical spines. Outline of exten- 123.8 mm, cw 122.3 mm), 41°39.6’S, 169°20.9’E, 1025 sor margin of walking legs generally even in adults, m, FV Arrow, A03/49/83, 7 Oct 1983. without several prominent spines protruding above Bay of Plenty: NIWA 6713, 1 juvenile male (cl 24.8 level of general spines [Australia]...... N. flindersi mm, pcl 13.7 mm, cw 11.6 mm), Otara Seamount, – Pereopod 2–4 coxae in both sexes covered with low, 36°56.81–57.09’S, 177°20.09–19.90’E, 1323–1346 m, blunt nodules or tubercles (occasionally subacute TAN0413/41, 10 Nov 2004; NIWA 6712, 1 indet. in juveniles). Outline of extensor margin of walk- juvenile (cl 13.7 mm, pcl 8.6 mm, cw 7.3 mm), Otara ing legs uneven in adults, with several prominent Seamount, 36°57.32–57.26’S, 177°20.93–20.67’E, spines protruding above level of general spines 1080–1081 m, TAN0413/29, 9 Nov 2004; NIWA 34911, [New Zealand]...... N. brodiei 1 male (pcl 97.4 mm, cw 96.1 mm), 37°05.80–03.40’S, 176°43.80–38.59’E, 1071–1049 m, Z10064, 27 Mar 2000; NIWA 34912, 1 male (pcl 112.7 mm, cw 106.2 mm), Neolithodes brodiei Dawson & Yaldwyn, 1970 Nukuhou Knoll, 37°16.0’S, 177°17.3’E, 1130 m, FV (Figs 43–48, Pl. 2A) Ocean Fresh, Z10063, 29 Mar 2000; NMNZ Cr11786, 1 ovigerous female (crushed), Bay of Plenty, 37°17.5’S, Neolithodes brodiei Dawson & Yaldwyn, 1970: 227–228 [type 176°58.67’E, 1000 m, crab pot, FV Savannah, coll. G. locality: E of Campbell Plateau, 50°58’S, 173°57’E, 832 m]. Gibbs, 8 Oct 2001. — Dawson & Yaldwyn, 1985: 70 [New Zealand occur- Hawkes Bay: NMNZ Cr8589, 1 male (cl 177.6 mm, rences]. — McLay, 1988: 36–38, fig. 3. — Dawson, 1989: pcl 155.5 mm, cw 149.9 mm), Ritchie Bank, 39°37’S, 3, frontispiece, 318. — Takeda in Amaoka et al., 1990: 178°25’E; NMNZ Cr4812, 1 ovigerous female (cl 129.9 361. — Webber, 1997: 81, figs 1–3, 5. — O’Driscoll et al., mm, pcl 105.8 mm, cw 100.1 mm), off Portland Island, 2003: 65. — Davie, 2002: 73 [New Zealand occurrences]. — Zaklan, 2002: 768, 785, 804 [New Zealand occurrences]. 39°43.0’S, 177°53.3’E, 1120 m, Orange Roughy trawl, — Batson, 2003: 136–137. — Webber & Naylor, 2004a: FV Arrow, A01/17/87, 22 Jun 1987; NIWA 34910, 79. — Thatje & Lörz, 2005: fig. 2B [note: figure caption 1 male (cl 137.0 mm, pcl 112.9 mm, cw 107.5 mm), transposed]. — Ahyong et al., 2007: 155. 39°46.59–45.76’S, 178°09.28–10.05’E, 1250–1200 m, NOT Neolithodes brodiei. — Thatje & Lörz, 2005: 335–336, fig. TAN0109/22, Z10873, 4 Jul 2001; NMNZ Cr3753, 1 2A. [= N. yaldwyni Ahyong & Dawson, 2006] [note: figure ovigerous female (cl 142.6 mm, pcl 127.4 mm, cw 111.8 caption transposed]. mm), 39°47.7–40.8’S, 177°58.4–00.6’E, 1160–1240 m, FV NOT Neolithodes brodiei. — Macpherson, 2001: 799 [= unde- Kalinovo, K01/22/81, 24 Nov 1981; NMNZ Cr3754, 1 scribed species]. male (pcl 146.0 mm, cw 144.3 mm), 39°47.7–40.8’S, NOT Neolithodes brodiei. — Dawson & Yaldwyn, 1985: 70. — Davie, 2002: 73 [Tasmania and Victoria]. — Zaklan, 177°58.4–00.6’E, 1160–1240 m, FV Kalinovo, K01/22/81, 2002: 768 [Australian occurrences]. — Poore, 2004: 268, 24 Nov 1981; NMNZ Cr3755, 1 ovigerous female (cl fig. 75. [= N. flindersi Ahyong, 2010]. 151.5 mm, pcl 123.9 mm, cw 110.0 mm), 39°47.7–40.8’S, 177°58.4’E–178°00.6’E, 1160–1240 m, FV Kalinovo, 24 Type material. Holotype: NIWA 735, ovigerous female Nov 1981; NIWA 25531, 1 male (pcl 48.8 mm, cw 44.0 (pcl 111.5 mm, cw 103.9 mm), Campbell Plateau, mm), 39°49.99’S, 177°39.00’E, 1000–1100 m, Z8371, 10 50°58.00’S, 173°56.99’E, 832 m, Globigerina ooze, small Aug 1995; NIWA 34909, 1 male (pcl 98.8 mm, cw 96.0 Agassiz trawl, HMNZ Endeavour, F135, 30 Jan 1965. mm), 39°49.99’S, 177°39.00’E, 1000–1100 m, Z8371, 10 Aug 1995; NMNZ Cr3751 P404, 1 male (pcl 134.0 mm, Other material examined. West Norfolk Ridge: NMNZ cw 126.9 mm), 40°47’S, 177°05’E, 1055–1180 m, FV Cr11150, 1 male (pcl 111.4, cw 101.0 mm), 34°34.26– Kalinovo, K01/147/81, 18 Dec 1981. 33.37’S, 168°56.53–54.76’E, 1013–1350 m, TAN0308/156, Wairarapa coast: NMNZ Cr3793, 1 male (cl 130.0 mm, 4 Jun 2003. pcl 101.5 mm, cw 95.0 mm), 30.5 nautical miles [56 km] Challenger Plateau: NIWA 3919, 1 damaged juvenile off Cape Turnagain, 1006–1189 m, FV Twofold Bay, coll. female, 37°27.30–28.80’S, 168°14.29–29.89’E, 953–992 A.D. Carruthers, 28 Dec 1984; NMNZ Cr3840–3843, 3 m, MFish SOP 1630/23, FV Baldur, coll. S. Smith, 24 males (pcl 48.1–115.6 mm, cw 44.5–111.3 mm), 1 female Mar 2002; NMNZ Cr3747, 1 male (pcl 86.2 mm, cw (cl 123.4 mm, pcl 103.8 mm, cw 102.5 mm), off Cape

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 43. Neolithodes brodiei Dawson & Yaldwyn, 1970, male, pcl 115.6 mm, off Cape Palliser (NMNZ Cr3840). A, dorsal habitus. B, carapace, right lateral view.

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 44. Neolithodes brodiei Dawson & Yaldwyn, 1970, male, pcl 115.6 mm, off Cape Palliser (NMNZ Cr3840). A, right chela. B, left chela. C, left pereopod 4. D, anterior carapace, dorsal view. E, left pereopod 4 dactylus. F, right orbit and antenna. G, anterior carapace, right lateral view.

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 45. Neolithodes brodiei Dawson & Yaldwyn, 1970, male, pcl 115.6 mm, off Cape Palliser (NMNZ Cr3840). A, carapace. B, ventral surface and abdomen. C, posterior carapace and abdominal somite 2.

Palliser, Orange Roughy depth, Apr/May 1985; NMNZ 803–1120 m, RV James Cook, J18/7/84, 16–18 Oct 1984; Cr3791, 1 male (cl 172.0 mm, pcl 135.1 mm, cw 135.2 NMNZ Cr3837, 1 male (pcl 40.2 mm, cw 38.0 mm), off mm), off Cape Palliser, 40°00’S–41°30’S, 176°00–31’E, Castle Point, 40°51.2–49.1’S, 176°55.9–55.7’E, 1030–1060

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ m, FV Kalinovo, K1/40/81, 25 Nov 1981; NMNZ 60581, 1 male (cl 97.0 mm, pcl 76.9 mm, cw 70.9 mm), Cr3838, 1 male (cl 83.7 mm, pcl 53.0 mm, cw 48.7 44°08.20–08.19’S, 178°47.90–46.61’E, 921–922 m, FV mm), off Castle Point, 40°51.5–53.6’S, 176°57.9–56.3’E, San Waitaki, SWA0901/88, 17 Nov 2009; NIWA 34895, 1125–1150 m, FV Kalinovo, K1/40/81, 28 Nov 1981; 1 male (cl 79.0 mm, pcl 57.7 mm, 51.8 mm), 44°12.39– NMNZ Cr3789, 1 female (cl 142.1 mm, pcl 122.3 mm, cw 12.99’S, 179°03.54–0.46’E, 959–961 m, TAN9812/15, 108.1 mm), off Cape Palliser, 41°00–30’S, 176°00–31’E, Z9468, 2 Oct 1998; NIWA 60580, 1 female (pcl 64.6 803–1107 m, RV James Cook, J18/-/84, 16–18 Oct 1984; mm, cw 59.3 mm), 44°22.83–22.66’S, 179°54.68–56.27’E, NMNZ Cr3745, 1 male (pcl 129.3 mm, cw 126.1 mm), 952–957 m, FV San Waitaki, SWA0901/82, 16 Nov off Wairarapa, 41°20.4’S, 176°16.3’E, 1070–1080 m, 2009; NIWA 34613, 1 female (pcl 70.2 mm, cw 64.2 RV James Cook, J6/9/84, 2 Apr 1984; NMNZ Cr3839, mm), 44°25.26–26.28’S, 174°59.30’W–175°00.75’W, 1 juvenile female (pcl 32.3 mm), E of Cape Palliser, 913–936 m, TAN0709/115, 22 Jul 2007; NIWA 60578, 41°29.7–23.8’S, 176°08.4–12.1’E, 1103–1071 m, RV James 1 male (pcl 149.4 mm, cw 149.2 mm), 44°32.78–33.62’S, Cook, J18/4/84, 16 Oct 1984; NMNZ Cr3752, 1 damaged 175°23.73–23.77’W, 939–985 m, FV San Waitaki, male (pcl 116.2 mm), SE of Castle Point, 41°13.7–15.4’S, SWA0901/61, 13 Nov 2009; NIWA 60556, 1 ovigerous 176°36.1–30.9’E, 1040–1100 m, FV Kalinovo, K01/35/81, female (cl 151.6 mm, pcl 122.4 mm, cw 113.2 mm), 27 Nov 1981; NMNZ Cr3775, 1 female (pcl 105.0 mm, 44°34.38–35.02’S, 175°20.27–20.28’W, 1070–1106 m, FV cw 95.4 mm), off Wairarapa, 41°21.5’S, 176°20.9’E, San Waitaki, SWA0901/060, 13 Nov 2009; NIWA 34896, 1073–1112 m, RV James Cook, J6/14/84, 3 Apr 1984; 1 female (pcl 31.5 mm, cw 28.5 mm), 44°35.28–34.90’S, NMNZ Cr3776, 1 male (pcl 66.7 mm, cw 61.4 mm), off 177°27.43–29.03’W, 852–858 m, TAN9812/75, Z9461, Wairarapa coast, 41°23.6’S, 176°21.7’E, 995–1082 m, 21 Oct 1998; NMNZ Cr3772, 1 female (pcl 95.3 mm, RV James Cook, J6/13/84, 3 Apr 1984; NMNZ Cr3777, cw 96.1 mm), off Canterbury, 44°45.1’S, 173°05.2’E, 1 male (cl 146.2 mm, pcl 119.6 mm, cw 118.8 mm), off 922–929 m, RV James Cook, J10/16/84, 10 Jun 1984; Wairarapa coast, 41°23.6’S, 176°21.7’E, 995–1082 m, NMNZ Cr9211, 1 juvenile female (cl 52.6 mm, pcl 35.4 RV James Cook, J6/13/84, 3 Apr 1984; NMNZ Cr3774, mm, cw 32.9 mm), 44°50.6’S, 175°29.4’E, 1044–1050 1 male (cl 170.6 mm, pcl 141.5 mm, cw 141.3 mm), m, RV James Cook, JCO90/015/5, 20 Sep 1990; NIWA, off Wairarapa, 41°27.5’S, 176°02.4’E, 1007–1075 m, 1 male (pcl 148.6 mm, cw 151.2 mm), Chatham Rise, RV James Cook, J6/7/84, 2 Apr 1984; NMNZ Cr3792, Tow 123, NEB. 1 ovigerous female (pcl 145.1 mm, cw 138.1 mm), Off West Coast: NMNZ Cr3786, 1 ovigerous female off Wairarapa coast, 41°28.4–26.1’S, 176°05.2–08.2’E, (cl 178.1 mm, pcl 153.0 mm, cw 145.6 mm), 43°15.3’S, 1121–1120 m, RV James Cook, J18/3/84, 3 16 Oct 1984; 168°27.2’E, 1027–1041 m, RV James Cook, J16/10/83, NMNZ Cr11570, 1 male (pcl 143.0 mm, cw 143.5 mm), 13 Dec 1983; off Wairarapa coast, 700 m, coll. D. Wylie, 17 Oct 1986; Louisville Ridge: NIWA 34893, 1 male (pcl 62.9 mm, NMNZ Cr11621, 1 male (pcl 133.2 mm, cw 133.1 mm), cw 58.3 mm), southern Louisville Ridge, 45°33.99’S, 2 females (cl 149.8–159.5 mm, pcl 127.6–132.1 mm, cw 156°40.00’W, 870–900 m, Z8370, FV Peterson, Sep 117.4–120.0 mm), off Wairarapa coast, 700 m, coll. D. 1995. Wylie, 13 Oct 1983. Campbell Plateau: NMNZ Cr3779, 1 male (pcl 73.9 Chatham Rise: NMNZ Cr7353, 1 male (pcl 145.9 mm, mm, cw 66.8 mm), Campbell Plateau (Pukaki Rise cw 142.4 mm), NW of Chatham Islands, 42°42.0–41.7’S, or near Campbell Island), RV Shinkai Maru, Stn 241; 178°01.0–03.7’W, 1025–1055 m, COR/223/89, 13 Aug NMNZ Cr3778, 1 parasitised female (cl 128.8 mm, pcl 1989; NIWA 33773, 1 ovigerous female (pcl 88.5 mm, 113.7 mm, cw 106.1 mm), Campbell Plateau (Pukaki cw 85.2 mm), 42°49.44–49.38’S, 175°29.87–31.93’W, Rise or near Campbell Island), RV Shinkai Maru, Stn 1105–1095 m, TAN0709/24, 9 Jul 2007; NMNZ Cr3758– 241; NMNZ Cr3750, 1 male (pcl 139.4 mm, cw 130.3 3759, 2 males (pcl 107.9–117.1 mm, cw 104.8–109.6 mm), 47°27’S, 169°26’E, 500 m, FV Wessermunde, mm), NE of Chathams, 42°55.4’S, 174°31.9’W, 900 WO 2B/165/79, 2 Mar 1979; NIWA 34894, 1 male m, FV Kalton, KTN61/82; NIWA 34588, 1 female (pcl (pcl 76.8 mm, cw 70.7 mm), 47°57.30’S–48°00.19’S, 99.0 mm, cw 92.5 mm), 42°55.62’S, 174°42.01–39.93’W, 169°55.83–57.01’E, 910 m, TAN0012/79, Z10849, 15 1076–1074 m, TAN0709/86, 17 Jul 2007; NIWA 34923, 1 Dec 2000; NIWA 60552, 1 male (pcl 80.5 mm, cw 73.6 ovigerous female (cl 145.0 mm, pcl 120.9 mm, cw 113.8 mm), 1 ovigerous female (pcl 104.7 mm, cw 98.3 mm), mm), NW Chatham Rise off Mernoo Bank, TAN0709, E of Pukaki Rise, 48°36.79’S, 175°15.03’E, 820–815 m, no specific station; NMNZ Cr14893, 1 juvenile male TAN0911/15, 1 Dec 2009; NIWA 44075, 1 male (pcl (pcl 34.5 mm, cw 30.1 mm), E of Chatham Islands, 109.8 mm, cw 102.5 mm), Pukaki Rise, 50°02–03’S, 43°01.06’S–42°59.62’S, 174°33.46–37.20’W, 979–988 m, 174°41–45’E, 855–975 m, with Briarosaccus callosus TAN9206/153, 6 Jul 1992; NIWA 41608, 1 female (pcl (Rhizocephala), SOP TRIP2468/70, 28 Aug 2007; 81.5 mm, cw 74.1 mm), 1 ovigerous female (pcl 97.3 NIWA 60554, 2 ovigerous females (pcl 104.1–111.7 mm, cw 94.6 mm), 44°05.49–06.33’S, 178°32.04–32.10’E, mm, cw 99.1–102.5 mm), SE of Pukaki Rise, 50°15.14’S, 952–982 m, TAN9812, Z9386, 30 Sep 1998; NIWA 174°05.49’E, 772–776 m, TAN0911/19, 2 Dec 2009;

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 46. Neolithodes brodiei Dawson & Yaldwyn, 1970, female, pcl 103.8 mm, off Cape Palliser (NMNZ Cr3841). A, dorsal habitus. B, ventral surface and abdomen.

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ NIWA 34892, 1 male (pcl 66.2 mm, cw 61.0 mm), beyond antennal peduncle by less than one-quarter Auckland Islands, 51°38.80’S, 164°35.28’E, 968–973 m, distal antennular peduncle article. TAN0306/8, 14 Apr 2003. Antenna: Basal antennal article with small anterola- New Zealand, no specific locality: NMNZ Cr3831, 1 teral spine; outer margin of article 2 with cluster of 4–7 male (cl 188.0 mm, pcl 165.5 mm, cw 161.6 mm), north- small spines basally and long slender spine that reaches ern New Zealand, FV Wanaka, 1984; NMNZ Cr3833, 1 to end of article 4; article 3 with sharp to angular inner male (cl 83.8 mm, pcl 55.5 mm, cw 50.6 mm), northern distal tooth; scaphocerite minute, often with bifid or New Zealand waters, FV Wanaka, 1984; NMNZ Cr3832, trifid apex, shorter than article 4; article 4 with mesial 1 female (pcl 121.4 mm, cw 112.1 mm), northern New and lateral distal spinule and occasionally 1–3 other Zealand waters, FV Wanaka, 1984; NMNZ Cr3835, 1 minute spinules, about half as long as article 5; article male (cl 89.4 mm, pcl 75.1 mm, cw 68.8 mm), New 5 unarmed or with 1 or 2 minute acute granules. Zealand. Abdomen: Ornamentation similar in both sexes. Somite 2 densely spinose, with about 10 long spines Diagnosis. Carapace with numerous small secondary and numerous smaller spines on median plate; sub- spinules on the carapace and pereopods in addition to median and marginal plates densely spinose, spines the major spines. Cheliped dactyli with convex dorsal largest laterally; surface and margins of remaining margins, rounded in cross section. Antennal peduncle somites and telson multispinose. with few, small, scattered granules or minute spinules, Pereopod 1 (chelipeds): Dimorphic but spination simi- not distinctly spinose. Walking leg meri compressed, lar. Entire surface of both chelipeds, except for distal flattened; extensor margin with several upright major half or two-thirds of fingers, covered with numerous spines protruding above level of secondary spines; minute secondary spines in addition to major spines. pereopod 4 dactylus covered with spinules over sur- Coxa ventral surface with blunt tubercles and tufts faces of proximal half; ventral surfaces of coxae with of setae; ischiobasis with 4 or 5 stout ventral spines. blunt tubercles or granules. Merus inner margin with stout subdistal spine; ventral margin with two rows of stout spines, mesial row with Description. Carapace: Pyriform, 0.99–1.14 times longer 2 spines, lateral row with 3 or 4 spines; dorsal and than wide; dorsal surface armed with scattered, longer lateral surface spinose, spines largest distally. Carpus spines amongst dense, uniform covering of granules with prominent spines on dorsal and lateral surfaces, or minute secondary spinules. Gastric region bearing with 3 or 4 irregular rows of 3–5 spines. Palm of both 6 large spines forming transverse hexagon; with 2 chelipeds in both sexes with similar ornamentation; all near transverse rows of 3 smaller spines, first across surfaces spinose, though less pronounced mesially and centre of hexagon and second slightly anterior to ventrally; dorsal margin with 2 irregular rows of about hexagon. Hepatic spine strong, directed anterolater- 3 or 4 conical spines; midlateral surface with 2 rows ally; margin between outer orbital spine and hepatic of 4 or 5 spines of similar size to dorsal row; ventral spine with 1 or 2 spines. Branchial regions with 8–12 surface irregularly spinose. large dorsal spines in addition to scattered smaller Major cheliped 1.19–1.31 pcl (male), 1.16–1.23 (fe- secondary spines; margins with 12–16 major spines in male); upper palm length 1.07–1.11 times height (male), addition to numerous minute spines. Cardiac region 1.05–1.15 (female); occlusal margins of fingers corneous with 4 equal spines of similar size to gastric spines and for distal third to half, proximally with 3 calcareous 1 or 2 smaller spines near intestinal region. Intestinal nodules; dactylus dorsal margin broadly convex, with region with 2 upright spines in transverse row and 2 rows of tufts of golden setae and proximal cluster of or 3 smaller posterior directed spines. Pterygostomian 5–7 small spines, 1.29–1.35 times longer than dorsal region spinose or tuberculate, with small, anterior, margin of palm (male), 1.33–1.35 (female). submarginal spine. Minor cheliped 1.19–1.29 pcl (male), 1.16–1.20 pcl Rostrum 0.12–0.28 pcl; median spine inclined dor- (female); upper palm length 1.13–1.28 times height sally, with pair of divergent dorsal spines and smaller (male), 1.13–1.18 (female); occlusal margin corneous pair of basal spines; ventral proximal surface with in distal half, proximally crenulate; dactylus dorsal cluster of spinules. Posterior orbital margin concave margin broadly convex, with rows of tufts of golden in lateral view, margin usually unarmed, occasionally setae and proximal cluster of 5–8 small spines proxi- with 1–4 minute spinules; outer orbital spine slender, mally, 1.70–1.83 times longer than dorsal margin of reaching to but not beyond cornea (when eyes directed palm (male), 1.65–1.78 (female). forwards). Anterolateral spine shorter than outer or- Pereopods 2–4 (walking legs 1–3): Similar, slender, bital spine. elongate; ornamentation similar in both sexes; seg- Ocular peduncle: Longer than cornea; with scattered ments spinose, surface between major spines densely dorsal granules or spinules. covered with numerous minute secondary spinules or Antennule: Peduncle unarmed, reaching anteriorly granules. Pereopod 4 longest. Coxae with well-spaced,

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 47. Neolithodes brodiei Dawson & Yaldwyn, 1970. A, juvenile female, pcl 35.4, Chatham Rise (NMNZ Cr9211). B, juvenile male, pcl 48.1 (NMNZ Cr3843). C, male, pcl 86.2, Challenger Plateau (NMNZ Cr3747). D, ovigerous female, pcl 105.8 mm, Hawkes Bay (NMNZ Cr4812). E–F, ovigerous female holotype, pcl 111.5 mm, dorsal and ventral view, Campbell Plateau (NIWA 735).

blunt, rounded to flat tubercles, distal margins crenu- extensor margin with 6–9 major spines in addition to late to bluntly dentate. Ischiobasis with 3 or 4 stout paired distal spines; dorsal surface with row of 5–7 spines around distal margins and numerous smaller major spines of similar size to extensor spines; flexor ventral spinules. Merus dorsoventrally compressed, margin with 2 rows of 4–7 major spines; major spines flattened ovate in cross section, longer than propodus; on surface of merus distinctly standing out from

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 48. Geographic distributions of Neolithodes brodiei Dawson & Yaldwyn, 1970 (○) and N. bronwynae sp. nov. (●). Inset: bathymetric distribution of N. brodiei.

secondary spinules. Carpus slightly longer than half (female). Carpus 0.50–0.53 merus length (male), 0.55– merus length; with 5–7 spines on extensor margin and 0.59 (female). Propodus 0.85–0.87 merus length (male), 3 or 4 other major spines distributed on carpal surface, 0.92–0.93 (female); length:height ratio 8.52–9.14 (male), distal and second proximal spines longest, about twice 7.72–8.50 (female). Dactylus 0.61–0.68 propodus length as long as other spines. Propodus dorsoventrally com- (male), 0.59–0.60 (female). pressed; with 5–9 spines on extensor margin and 3–5 Pereopod 4 length 2.56–2.94 pcl (male), 2.36–2.62 similar spines on dorsal surface; flexor margin with pcl (female). Merus 0.84–0.99 pcl (male), 0.75–0.80 pcl numerous smaller spines. Dactylus curved, rounded (female); length:height ratio 4.54–5.06 (male), 4.26–4.31 in cross section, with 4 or 5 small proximal spines and (female). Carpus 0.52–0.55 merus length (male), 0.56– numerous secondary spines covering proximal half; 0.58 (female). Propodus 0.89–0.93 merus length (male), apex corneous. 0.98–1.03 (female); length:height ratio 8.76–9.14 (male), Pereopod 2 length 2.21–2.41 pcl (male), 2.01–2.09 8.03–9.33 (female). Dactylus 0.61–0.66 propodus length pcl (female). Merus 0.77–0.86 pcl (male), 0.66–0.68 pcl (male), 0.59–0.63 (female). (female); length:height ratio 4.41–5.04 (male), 3.80–4.05 (female). Carpus 0.51–0.60 merus length (male), 0.56– Colour in life. Deep-red overall. 0.63 (female). Propodus 0.83–088 merus length (male), 0.91–0.92 (female); length:height ratio 8.00–8.55 (male), Remarks. Neolithodes brodiei Dawson & Yaldwyn, 1970 6.43–7.71 (female). Dactylus 0.60–0.64 propodus length belongs to the group within the genus characterised (male), 0.62–0.63 (female). by bearing numerous secondary spinules on the cara- Pereopod 3 length 2.48–2.77 pcl (male), 2.30–2.43 pace and pereopods in addition to the major spines; pcl (female). Merus 0.85–0.99 pcl (male), 0.76–0.77 pcl convex dorsal margins of the cheliped dactylus; (female); length:height ratio 4.60–4.92 (male), 4.11–4.38 and compressed, flattened meri of the walking legs:

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ N. brodiei Dawson & Yaldwyn, 1970, N. nipponensis N. yaldwyni Ahyong & Dawson, 2006; southern Aus- Sakai, 1971, and N. flindersi Ahyong, 2010. Of these, tralian and Vanuatu records (Davie 2002; Poore 2004; N. brodiei most closely resembles N. flindersi Ahyong, Macpherson 2001) represent N. flindersi Ahyong, 2010, 2010 from southeastern Australia, previously misiden- and an undescribed species, respectively. Neolithodes tified as N. brodiei from that region. brodiei is not known from depths much beyond 1200 Neolithodes brodiei is best distinguished from m, so its distribution appears to be largely dictated by N. flindersi by patterns of spination: the ventral surfaces the extent of contiguous continental shelf and slope of the coxae of the walking legs in N. brodiei are covered habitat in the New Zealand region. in low, blunt tubercles rather than short, conical spines The only other species of Neolithodes known from (in males and juvenile female N. flindersi); the spinula- New Zealand, N. bronwynae sp. nov., markedly dif- tion covering the surfaces of the walking leg dactyli fers from N. brodiei by the very long dorsal spines in of N. brodiei usually extends only slightly beyond the adults, sparse rather than dense covering of secondary proximal half, rather than beyond the distal quarter; spinules on the carapace and pereopods, and subcylin- and in N. brodiei, the extensor margins of the pereopod dical rather than flattened meri of the walking legs. 2–4 meri are lined with short, closely-spaced spines Neolithodes brodiei and L. aotearoa sp. nov. are the that are interspersed by 6–9 distinctly longer spines, two largest species of lithodid in New Zealand waters, whereas in N. flindersi and N. nipponensis, the meral both being subject to fishery management under the extensor spines are of similar size. Neolithodes brodiei Quota Management System implemented by the New also differs from N. flindersi in more subtle features: Zealand Ministry of Fisheries. the rostrum in specimens exceeding 80 mm pcl is Batson (2003) reported the association between proportionally longer (0.12–0.28 versus 0.10–0.19 pcl); N. brodiei and the liparid fish,Careproctus sp., which in- the pereopod 4 propodus of N. brodiei is generally less cubates its eggs in the branchial chambers of the crab. slender than in N. flindersi (usually less than 10 times as long as high, with the merus longer than the propodus; Distribution. New Zealand and adjacent waters, from versus usually 10 times as long as high or greater, and the southern Norfolk Ridge to the Campbell Plateau, longer than the merus). Note that specimens with and the southern Louisville Ridge; 500–1240 m, usually regenerating limbs may be difficult to distinguish us- from 950–1150 m. ing pereopod 4 characters. Overall dorsal spination of juvenile N. brodiei is similar to juvenile N. flindersi and N. nipponensis but, as in adults, can be distinguished Neolithodes bronwynae sp. nov. by the ornamentation of the pereopod 2–4 coxae. (Figs 48–52, Pl. 2B) The strong morphological similarities and adjacent, but discrete, distributions suggest that N. brodiei and Neolithodes vinogradovi. — ?Macpherson, 1990: 218, fig. 2a [not N. flindersi might be sibling species. Neolithodes vinogradovi Macpherson, 1988b]. As with other lithodids, rostral and general spine length in N. brodiei is inversely proportional to body Type material. Holotype: NIWA 49026, male (cl 171.4 size, and the walking legs of females are proportionally mm, pcl 123.3 mm, cw 114.5 mm), Whakatane shorter than those of males. The median rostral spine Seamount, 36°47.71–47.28’S, 177°25.53–25.65’E, 1515– reaches up to 0.6 pcl in juveniles smaller than about 1530 m, TAN0413/19, 8 Nov 2004. 50 mm pcl, falling to about 0.1–0.3 pcl in specimens Paratype: NMNZ Cr11147, 1 male (cl 220.0 mm, pcl exceeding about 80 mm pcl. Bluntness of the tuber- 170.0 mm, cw 155.9 mm), Lord Howe Rise, 32°03.97– cles on the ventral surface of the pereopod 2–4 coxae 02.26’S, 159°52.80–51.10’E, 1934–1920 m, NORFANZ, also varies allometrically, being flattest and bluntest TAN0308/071, 24 May 2003. in adults. Occasionally, the tubercles of the coxae of pereopods 3–4 may be slightly pointed in juve- Diagnosis. Carapace dorsal surface with long, slender niles, approaching the condition of N. flindersi. The spines and scattered, widely separated granules or smallest ovigerous female measures 88.5 mm pcl minute secondary spines; longest spine exceeding 0.2 (NIWA 33773). pcl. Rostrum 0.3–0.4 pcl; ventral surface smooth. Poste- Neolithodes brodiei is the most common and wide- rior orbital margin near vertical; outer orbital spine not spread lithodid in New Zealand waters, ranging from reaching beyond cornea. Outer spine of antennal article the southern Norfolk Ridge south to the Campbell 2 reaching beyond article 4; scaphocerite reduced to Plateau, and east to the Louisville Ridge (156°W; out- a short, acute point, shorter than antennal article 4. side of the New Zealand Exclusive Economic Zone). Major cheliped palm with prominent dorsal and lateral Reports of N. brodiei from outside of the New Zealand spines; dactylus with convex dorsal margin, longer region are based on other species: Antarctic records than dorsal margin of palm. Walking legs spinose, sur- of N. brodiei (see Thatje & Lörz 2005) are referable to face between major spines finely granulate, secondary

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 49. Neolithodes bronwynae sp. nov., male holotype, cl 171.4 mm, pcl 123.3 mm, cw 114.5 mm, Whakatane Seamount (NIWA 49026). A, dorsal habitus. B, carapace, right lateral view.

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 50. Neolithodes bronwynae sp. nov., male holotype, cl 171.4 mm, pcl 123.3 mm, cw 114.5 mm, Whakatane Seamount (NIWA 49026). A, carapace. B, ventral surface and abdomen. C, right antenna. D, posterior carapace and abdominal somite 2.

spines absent or sparsely distributed; merus ovate in rows of 3 and 4 smaller spines, first across centre of cross section, shorter than or subequal to pcl. hexagon and second slightly anterior to hexagon. Hepatic spine strong, anterodorsally directed; margin Description. Carapace: Pyriform, 1.07–1.09 times longer between outer orbital spine and hepatic spine with 2 than wide; dorsal surface armed with long, slender spines. Branchial regions with 6 long dorsal spines in spines and scattered, widely separated granules or addition to scattered, minute, secondary spines; mar- minute secondary spines; longest spine 0.2–0.4 pcl (0.37 gins with 10 or 11 long spines in addition to several pcl in holotype). Gastric region bearing 6 long spines markedly shorter spines. Cardiac region with 4 long, forming transverse hexagon; with 2 near transverse equal spines of similar size to posterior gastric spines;

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 51. Neolithodes bronwynae sp. nov., male holotype, cl 171.4 mm, pcl 123.3 mm, cw 114.5 mm, Whakatane Seamount (NIWA 49026). A, right chela. B, left chela. C, right pereopod 4 dactylus and propodus. D, right pereopod 4 merus, distal end. E, anterior carapace, right lateral view.

with single shorter spine near intestinal region. Intes- Rostrum 0.29–0.39 pcl; median spine straight, tinal region with 2 long upright spines in transverse angled dorsally, with pair of long, divergent dorsal row and 2, much shorter, posterior directed spines. spines; ventral surface smooth. Posterior orbital margin Pterygostomian region spinose or tuberculate, with shallowly concave to almost straight in lateral view; stout, conical subdistal spine. outer orbital spine slender, reaching to but not beyond

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ cornea (when eyes directed forwards). Anterolateral granulate; length increasing posteriorly, pereopod 4 spine subequal to or shorter than outer orbital spine. longest. Distal margins of coxae with short triangular Ocular peduncle: Longer than cornea; with 5 dorsal spines. Ischiobasis with 4 distal spines (dorsal 2 short- granules or spinules. est) and 2 or 3 smaller ventral spines. Merus ovate in Antennule: Peduncle unarmed, reaching anteriorly cross section, shorter than carapace; extensor margin beyond antennal peduncle by half length of distal with 4 or 5 prominent spines increasing in length dis- antennular peduncle article. tally and intervening shorter spines, in addition to long, Antenna: Basal antennal article with small anterola- paired distal spines; dorsal surface with row of 5 spines teral spine; outer margin of article 2 with small basal of similar length to proximal extensor spines (except on spine and long slender spine that reaches proximal pereopod 4 merus with spine near midlength as long quarter to third of article 5; article 3 with angular inner as cardiac carapace spines); flexor margin with 2 rows distal tooth; scaphocerite minute, shorter than article of 5–7 spines (longest on pereopod 4). Carpus slightly 4, apex sharp; article 4 unarmed, about half length of longer than half merus length; extensor margin with article 5. 5 or 6 spines, distal and second proximal spines long- Abdomen: Somite 2 with 8–10 long spines of similar est, about twice length of other spines; dorsal surface length to cardiac spines, and several much shorter with 4 or 5 spines, distal and proximal spine longest. spines on median plate; submedian plate with 4 or Propodus ovate to subcircular in cross section; with 8 5 long spines near posterior margin and 5–7 shorter or 9 spines on extensor margin and 9–11 similar spines spines on surface; lateral border of marginal plates with on dorsal surface; flexor margin with 6–10 smaller 7 or 8 low, graded teeth proximally and 3 or 4 slender spines. Dactylus curved, rounded in cross section, with spines distally; surface and margins of remaining 4 small proximal spines and several scattered spinules somites multispinose or nodulose; telson spinose. on proximal surfaces; apex corneous. Pereopod 1 (chelipeds): Dimorphic but spination simi- Pereopod 2 length 2.51–2.52 pcl (male). Merus lar; surface of segments minutely granular. Coxa with 0.86–0.91 pcl (male); length:height ratio 6.41–6.59 blunt tubercles and tufts of setae, unarmed; ischiobasis (male). Carpus 0.51–0.52 merus length (male). Pro- with 5 or 6 stout ventral spines. Merus outer surface podus 0.86–0.89 merus length (male); length:height spinose, longest spine distally; ventral margin with 2 ratio 10.10–10.18 (male). Dactylus 0.66–0.50 propodus rows stout spines, mesial row with 2 spines, lateral row length (male). with 3 spines; dorsal and lateral surface spinose, spines Pereopod 3 length 2.70–2.73 pcl (male). Merus 0.92– largest distally. Carpus dorsal margin with irregular 0.98 pcl (male); length:height ratio 6.52–6.31 (male). row of 4 or 5 slender spines; lateral margin with 2 ir- Carpus 0.52–0.50 merus length (male). Propodus regular rows of 3 or 4 slender spines of similar size to 0.91–0.85 merus length (male); length:height ratio 10.29 dorsal row; ventrally with small, scattered, acute tuber- (male). Dactylus 0.64–0.52 propodus length (male). cles. Palm prominently spinose on dorsal, lateral and Pereopod 4 length 2.81–2.79 pcl (male). Merus ventral surfaces, inner surface with acute tubercles and 0.91–0.99 pcl (male); length:height ratio 6.84–6.62 small spines; dorsal margin with 2 irregular rows of 3 (male). Carpus 0.54–0.52 merus length (male). Pro- or 4 conical spines; midlateral surface with 2 irregular podus 0.96–0.91 merus length (male); length:height rows of 4 or 5 spines of similar size to dorsal rows; ratio 10.14–11.10 (male). Dactylus 0.66–0.51 propodus ventral surface with 2 rows of 4 or 5 spines, smaller length (male). than lateral and dorsal spines. Major cheliped 1.44–1.40 pcl (male); upper palm Colour in life. Deep-red overall (Pl. 2B). length 1.27–1.21 times height (male); occlusal margins corneous for slightly less than distal half, proximally Etymology. Named for Bronwyn Ahyong. with 3 calcareous nodules; dactylus dorsal margin broadly convex, with rows of tufts of golden setae and Remarks. On the basis of the long dorsal spines retained proximal cluster of 4–7 small spines, 1.29–1.07 times in adults, Neolithodes bronwynae sp. nov. resembles longer than dorsal margin of palm (male). N. vinogradovi Macpherson, 1988b [type locality: south- Minor cheliped 1.45–1.34 pcl (male); upper palm east Indian Ocean] and N. duhameli Macpherson, 2004 length 1.39–1.35 times height (male); occlusal margins [type locality: Crozet Islands]. Neolithodes bronwynae corneous for slightly more than distal half, proximally differs from N. duhameli in its proportionally longer, crenulate; dactylus dorsal margin broadly convex, with more slender walking legs that lack a dense covering of rows of tufts of golden setae and proximal cluster of secondary spinules and acute tubercles. The third walk- 4–6 small spines, 1.70–1.51 times longer than dorsal ing leg in N. bronwynae is 2.8 pcl versus 1.9–2.3 pcl in margin of palm (male). N. duhameli, with the merus about 0.9 rather than 0.7 Pereopods 2–4 (walking legs 1–3): Similar; segments pcl., and the propodi about 10 rather than 5–6 times prominently spinose, surface between major spines longer than high. The dense secondary spinulation

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 52. Neolithodes bronwynae sp. nov., male paratype, cl 220.0 mm, pcl 170.0 mm, cw 155.9 mm, Lord Howe Rise (NMNZ Cr11147). A, dorsal habitus. B, carapace.

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ of the walking legs in N. duhameli is replaced in 1 male (cl 52.4 mm, pcl 40.3 mm, cw 35.6 mm), 1 female N. bronwynae by a rather even granulation (Fig. 51D). (pcl 56.1 mm, cw 48.4 mm), off W coast of Tasmania, Although the secondary ornamentation of the walking 40°45’S, 143°31’E, 950 m, FRV Challenger, coll. M. Wil- legs differs between N. bronwynae and N. duhameli, both son, 16 Dec 1981; TM G2975, 1 female (pcl 55.8 mm, cw have similar secondary spinulation on the carapace, 49.9 mm), off W coast of Tasmania, 40°45’S, 143°30’E, distinguishing them both from N. vingradovi, which 860–1060 m, FRV Challenger, coll. M. Wilson, 9 Jan 1982; bears very few, scattered secondary spinules. Neolith- TM G3480, 1 juvenile male (cl 41.1 mm, pcl 27.6 mm, odes bronwynae further differs from N. vinogradovi in cw 22.7 mm), E coast of Tasmania, 41°27’S, 148°40’E, its shorter walking legs with a granular rather than FRV Soela, SO4/86/27, coll. P. Last, 13 Jul 1986; NMV smooth surface between the major spines. The third J12942, 1 female (pcl 101.0 mm, cw 91.5 mm), off NW walking leg of Neolithodes bronwynae, in comparison to Tasmania, 43°45’S, 143°40’E to 41°12’S, 143°55’E, N. vinogradovi, measures 2.8 versus 3.5 pcl; the merus is 500–650 fm [915–1189.5 m], FV Derwent Venture, coll. shorter than or almost as long as pcl versus 1.2 pcl. G. Switzer, 4–8 May 1986. The holotype and paratype agree well, differing chiefly in size-related features: the dorsal spines of Other material examined. New South Wales: AM P35596, the latter are proportionally shorter, the granules on 1 female (cl ~76 mm, badly damaged), E of Broken Bay, the walking legs are slightly less distinct, and the dac- 33°31–28’S, 152°12–14’E, 1079–951 m, FRV Kapala, K83- tyli of the walking legs are proportionally shorter (0.5 15-03, 3 Nov 1983; AM P35604, 1 male (cl 161.6 mm, pcl versus 0.6–0.7 propodus length). The posterior mar- 135.5 mm, cw 132.9 mm), E of Brush Island, 35°38’S, gin and right branchial surface of the carapace of the 150°44’E, 887 m, FRV Kapala, K83-11-03, coll. K. Gra- paratype exhibits evidence of prior damage, perhaps ham, 8 Sep 1983; AM P81161, 1 male (cl 124.1 mm, pcl from moulting. 105.8 mm, cw 93.0 mm), off Bermagui, 36°10.8–15.0’S, Macpherson (1990) reported a large female (pcl 105 150°24.4–22.4’E, 457.5–503.3 m, FV Shelley H, coll. K. mm, cw 91 mm) from New Caledonia as N. vinogradovi Graham, 1 Sep 1999; AM P35598, 1 male (pcl 149.2 mm, that, apart from sexual dimorphism, differed from the cw 149.2 mm), E of Gabo Island, 37°37’S, 150°21’E, 960 male holotype in more numerous secondary spines. In m, FRV Kapala, K84-04-04, coll. K. Graham, 5 Apr 1984; view of the close geographic proximity and similar sec- AM P35603, 1 male (pcl 112.7 mm, cw 109.5 mm), E of ondary spination, the New Caledonian material might Brush Island, 35°31–28’S, 150°50–53’E, 942–978 m, FRV be referable to N. bronwynae. Neolithodes bronwynae is Kapala, K83-11-04, coll. K. Graham, 8 Sep 1983; AM readily distinguished from the only other New Zealand P35606, 1 male (cl 124.0 mm, pcl 110.3 mm, cw 101.1 congener, N. brodiei, by its long dorsal spines and the mm), E of Brush Island, 35°31’S, 150°50’E, 978 m, on granular, rather than densely spinose, surfaces between Chrysogorgia orientalis, FRV Kapala, K83-11-04, coll. K. the major spines of the walking legs. Graham, 8 Sep 1983. Victoria: AM P52067, 1 female (cl 146.9 mm, pcl 126.2 Distribution. Presently known from the Whakatane mm, cw 112.5 mm), E of Gabo Island, 37°42.1–39.4’S, Seamount (Bay of Plenty) and the Lord Howe Rise 150°16.6–17.6’E, 604–617 m, FRV Kapala, K97-01-02, near Lord Howe Island (Tasman Sea); possibly also coll. K. Graham, 16 Apr 1997; AM P35592, 1 female (pcl from New Caledonia; 1515–1920 m. 38.6 mm, cw 32.4 mm), E of Gabo Island, 37°43–41’S, 150°20’E, 960–978 m, FRV Kapala, K83-12-01, 26 Sep Neolithodes flindersi Ahyong, 2010 1983; SAM C6854, 1 male (cl 86.3 mm, pcl 75.3 mm, cw 68.8 mm), 65 nautical miles [120 km] SE of Portland, (Figs 53–58, Pl. 2C) 39°21.55–20.66’S, 142°40.54–39.75’E, demersal trawl, Neolithodes brodiei. — Dawson & Yaldwyn, 1985: 70. — Davie, RV Soela, coll. K. Gowlett-Holmes, 4 Mar 1989. 2002: 73 [Tasmania and Victoria]. — Zaklan, 2002: 768 South Australia: SAM C6855, 1 female (cl 110.0 mm, [Australian occurrences]. — Poore, 2004: 268, fig. 75. [Not pcl 96.2 mm, cw 89.0 mm), Great Australian Bight, ap- N. brodiei Dawson & Yaldwyn, 1970]. prox. 120 nautical miles [222 km] SW of Cape Adieu, Neolithodes flindersi Ahyong, 2010: 56–62, figs 1–3 [type local- 33°58’S, 131°22’E, 1000 m, trawled, FV Saxon Progress, ity: NW Tasmania, about 900 m]. coll. D. Wheenan, Nov 1989; SAM C6856, 1 male (pcl 83.9 mm, cw 78.4 mm), Great Australian Bight, ap- Type material. Holotype: NMV (ex NMNZ Cr011763), prox. 120 nautical miles [222 km] SSE of Eucla, 33°39’S, male (cl 183.3 mm, pcl 126.1 mm, cw 115.8 mm), NW 129°50’E, 984–1015 m, FV Longva III, coll. K. Gowlett- Tasmania, about 900 m, otter trawl, FV Margaret Holmes, 12 Dec 1989; SAM C6857, 1 female (cl 110.6 Philippa, coll. W. Nicholls, 1983. mm, pcl 99.23 mm, cw 91.3 mm), Great Australian Paratypes: NMNZ Cr011764, female (pcl 117.0 mm, Bight, approx. 135 nautical miles [250 km] SSW of Cape cw 107.3 mm), NW Tasmania, about 900 m, otter trawl, Adieu, 34°13’S, 131°30’E, 1122–1330 m, from black FV Margaret Philippa, coll. W. Nicholls, 1983; TM G2973, coral ‘A’, FV Longva III, coll. K. Gowlett-Holmes, 19

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 53. Neolithodes flindersi Ahyong, 2010, male holotype, cl 183.3 mm, pcl 126.1 mm, cw 115.8 mm, NW Tasmania (NMV, ex NMNZ Cr011763). A, dorsal habitus. B, carapace.

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 54. Neolithodes flindersiAhyong, 2010, male holotype, cl 183.3 mm, pcl 126.1 mm, cw 115.8 mm, NW Tasma- nia (NMV, ex NMNZ Cr011763). A, right chela. B, left chela. C, ventral surface and abdomen. D, right pereopods 2–3 coxae. E, posterior carapace and abdominal somite 2. F, carapace, right lateral view.

Dec 1989; TM G2977, 1 male (cl 75.4 mm, pcl 55.0 mm, Bight, approx. 120 nautical miles [222 km] SSW of Cape cw 48.8 mm), 20 miles [32 km] SW of Port MacDon- Adieu, 33°57’S, 131°20’E, 1004–1030 m, FV Longva III, nell, 38°19–23’S, 140°20–32’E, 800–1000 m, FV Tuna coll. K. Gowlett-Holmes, 30 Nov 1989; SAM C6356, 1 Endeavour, coll. R.M. Green, 5 Aug 1983; SAM C6355, male (cl 105.0 mm, pcl 92.8 mm, cw 82.0 mm), 1 female 1 female (pcl 97.3 mm, cw 96.1 mm), Great Australian (cl 81.0 mm, pcl 67.1 mm, cw 57.5 mm), Great Austral-

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 55. Neolithodes flindersi Ahyong, 2010. A–E, male holotype, cl 183.3 mm, pcl 126.1 mm, cw 115.8 mm, NW Tasmania (NMV, ex NMNZ Cr011763). F, male, pcl 55.0 mm, SW of Port MacDonnell (TM G2977). A, anterior carapace, dorsal view. B, right orbit and antenna. C, right pereopod 4. D, right pereopod 4 merus. E, right pereopod 4 dactylus. F, right pereopod 4 merus and carpus.

ian Bight, approx. 120 nautical miles [222 km] SSW of male (pcl 108.6 mm, cw 96.2 mm), 1 female (cl 93.0 mm, Cape Adieu, 34°01’S, 131°18’E, 994–1065 m, FV Longva pcl 82.3 mm, cw 74.1 mm), Great Australian Bight, ap- III, coll. K. Gowlett-Holmes, 14 Dec 1989; SAM C6360, 1 prox. 120 nautical miles [222 km] SSW of Cape Adieu,

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 56. Neolithodes flindersi Ahyong, 2010, female paratype, pcl 117.0 mm, cw 107.3 mm, NW Tasmania (NMNZ Cr11764). A, dorsal habitus. B, abdomen.

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ 33°59’S, 131°27’E, 1020 m, FV Longva III, stn 136, coll. K. smaller secondary spines; margins with 13–15 major Gowlett-Holmes, 3 Dec 1989; SAM C6359, 1 female (cl spines in addition to numerous minute spines. Cardiac 105.2 mm, pcl 94.1 mm, cw 87.4 mm), Great Australian region with 4 equal spines of similar size to gastric Bight, approx. 120 nautical miles [222 km] SSW of Cape spines and 1 or 2 smaller spines near intestinal region. Adieu, 33°57’S, 131°19’E, 1020–1055 m, FV Longva III, Intestinal region with 2 upright spines in transverse coll. K. Gowlett-Holmes, 22 Nov 1989; SAM C6365, 1 row and 2 or 3 smaller, posteriorly directed spines. male (cl 114.7 mm, pcl 99.4 mm, cw 92.3 mm), 32–35 Pterygostomian region spinose or tuberculate, with miles [51–56 km] off Beachport, 1333 m, trawl, FV Silent small, anterior, submarginal spine. Venture, coll. W. Rumball, 27–29 Nov 1987; SAM C6362, Rostrum 0.09–0.19 pcl; median spine inclined dor- 1 male (cl 144.9 mm, pcl 128.8 mm, cw 125.9 mm), Great sally, with pair of divergent dorsal spines and smaller Australian Bight, approx. 120 nautical miles [222 km] pair of basal spines; ventral proximal surface with SSW of Cape Adieu, 33°57’S, 131°20’E, 954–1016 m, cluster of spinules. Posterior orbital margin concave in FV Longva III, coll. K. Gowlett-Holmes, 30 Nov 1989; lateral view, occasionally with 2–4 spinules on lower SAM C6363, 1 male (pcl 154.1 mm, cw 151.5 mm), Great margin; outer orbital spine slender, not reaching cornea Australian Bight, approx. 120 nautical miles [222 km] (when eyes directed forwards). Anterolateral spine as SSW of Cape Adieu, 33°57’S, 131°22’E, 1012–1033 m, FV long as or shorter than outer orbital spine. Longva III, coll. K. Gowlett-Holmes, 30 Nov 1989; SAM Ocular peduncle: Longer than cornea; with scattered C6364, 1 ovigerous female (cl 165.3 mm, pcl 151.1 mm, dorsal granules or spinules. cw 143.8 mm), Great Australian Bight, approx. 95 nauti- Antennule: Peduncle unarmed, reaching anteriorly cal miles [176 km] W of Cape Wiles, 34°57’S, 133°20’E, beyond antennal peduncle by less than half distal an- 850–970 m, FV Longva III, coll. K. Gowlett-Holmes, 11 tennular peduncle article. Nov 1989; NMNZ Cr20314–20315 (ex SAM), 2 males (cl Antenna: Basal antennal article with small antero- 89.0–94.3 mm, pcl 75.3–77.0 mm, cw 70.7–74.8 mm), 30 lateral spine; outer margin of article 2 with or without nautical miles [56 km] SW of Cape Martin Lighthouse, cluster of small spinules basally and long slender spine 37°48.6’S, 139°30.99’E, 1000–1060 m, trawl, FV Silent that reaches to end of article 4; article 3 with 1 or 2 sharp Victory, K. Gowlett-Holmes et al., 15 Dec 1987; SAM to angular inner distal teeth; scaphocerite minute, often TC15379, 1 juvenile male (cl 31.9 mm, pcl 21.6 mm, with bifid or trifid apex, shorter than article 4; article 4 cw 18.4 mm), 31 nautical miles [57 km] SW of Cape with mesial and lateral distal spinule and occasionally 1 Martin lighthouse, Beachport, 37°48.61’S, 139°29.74’E, or 2 other minute spinules, about half as long as article 933–1098 m, FV Silent Victory, coll. K. Gowlett-Holmes 5; article 5 with scattered spinules or granules. et al., 16 Dec 1987. Abdomen: Ornamentation similar in both sexes. Somite 2 densely spinose, with about 10 long spines Diagnosis. Carapace with numerous small secondary and numerous smaller spines on median plate; sub- spinules on the carapace and pereopods in addition median and marginal plates densely spinose, spines to major spines. Antennal peduncle with few, small, largest laterally; surface and margins of remaining scattered granules or minute spinules, not distinctly somites and telson multispinose. spinose. Cheliped dactyli with convex dorsal margins, Pereopod 1 (chelipeds): Dimorphic but spination simi- rounded in cross section. Walking leg meri compressed, lar. Entire surface of both chelipeds, except for distal flattened ovate in cross section; larger spines on exten- half or two-thirds of fingers, covered with numerous sor margin not protruding above level of secondary minute secondary spines in addition to major spines. spines; dactyli of pereopod 4 covered with spinules Coxa ventral surface with blunt tubercles and tufts onto proximal three-quarters; ventral surfaces of coxae of setae; ischiobasis with 4 or 5 stout ventral spines. with short conical spines in males and juvenile females Merus inner margin with stout subdistal spine; ventral (obsolete in adult females). margin with two rows of stout spines, mesial row with 2 spines, lateral row with 3 or 4 spines; dorsal and Description. Carapace: Pyriform, 1.01–1.15 times longer lateral surface spinose, spines largest distally. Carpus than wide; dorsal surface armed with scattered major with prominent spines on dorsal and lateral surfaces, spines amongst dense, uniform covering of small sec- with 3 or 4 irregular rows of 3–5 spines. Palm of both ondary spinules. Gastric region bearing 6 large spines chelipeds in both sexes with similar ornamentation; all forming transverse hexagon; with 2 near transverse surfaces spinose, although less pronounced mesially rows of 3 smaller spines, first across centre of hexagon and ventrally; dorsal margin with 2 irregular rows of and second slightly anterior to hexagon. Hepatic spine about 3 or 4 conical spines; midlateral surface with anterolaterally directed; margin between outer orbital 2 rows of 4 or 5 spines of similar size to dorsal row; spine and hepatic spine with 2 or 3 larger spines in ad- ventral surface irregularly spinose. ditional to small secondary spines. Branchial regions Major cheliped 1.27–1.35 pcl (male), 1.13–1.14 (fe- with 8–10 larger dorsal spines in addition to scattered male); upper palm length 1.06–1.14 times height (male),

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 57. Neolithodes flindersi Ahyong, 2010. A, juvenile male paratype, E Tasmania, pcl 27.6 mm (TM G3480). B, male, pcl 55.0 mm, SW of Port MacDonnell (TM G2977).

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ 1.14–1.16 (female); occlusal margins of fingers corneous Pereopod 4 length 3.05–3.14 pcl (male), 2.41–2.87 for distal third to half, proximally with 3 calcareous pcl (female). Merus 0.86–0.94 pcl (male), 0.70–0.86 pcl nodules; dactylus dorsal margin broadly convex, with (female); length:height ratio 3.98–5.01 (male), 3.49–4.22 rows of tufts of golden setae and proximal cluster of (female). Carpus 0.56–0.57 merus length (male), 4–7 small spines, 1.23–1.33 times longer than dorsal 0.59–0.62 (female). Propodus 1.03–1.10 merus length margin of palm (male), 1.26–1.43 (female). (male), 1.08–1.14 (female); length:height ratio 9.67– Minor cheliped 1.24–1.32 pcl (male), 1.09–1.21 pcl 12.27 (male), 8.41–11.42 (female). Dactylus 0.58–0.60 (female); upper palm length 1.17–1.33 times height propodus length (male), 0.62–0.70 (female). (male), 1.17–1.31 (female); occlusal margin corneous in distal half, proximally crenulate; dactylus dorsal Colour in life. Deep-red overall (Pl. 2C). margin broadly convex, with rows of tufts of golden Remarks. Recognition of N. brodiei from Australia stems setae and proximal cluster of 4–6 small spines proxi- from Dawson & Yaldwyn’s (1985: 70) remark that the mally, 1.69–1.98 times longer than dorsal margin of species “occurs in appreciable numbers off south-east- palm (male), 1.65–1.86 (female). ern Australia” based on collections in the Australian Pereopods 2–4 (walking legs 1–3): Similar, slender, Museum. All of the Australian Museum specimens, elongate; ornamentation similar in both sexes; seg- however, as well as other Australian specimens iden- ments spinose, surface between major spines densely tified as N. brodiei in other collections, are referable to covered with small spines. Pereopod 4 longest. Coxae N. flindersi. Records of N. brodiei from Vanuatu and N. covered with small conical spinules in males, in fe- nipponensis from the Solomon Islands (Macpherson males becoming obsolete or reduced to blunt gran- 2001, 2003) may represent undescribed species under ules beneath overlap of abdomen (covered in conical study by P. Davie. spinules in juvenile females); distal margins crenulate Neolithodes flindersi most closely resembles N. brodiei to bluntly dentate. Ischiobasis with 3–5 stout spines Dawson & Yaldwyn, 1970, from New Zealand, and around distal margins and numerous, smaller, ventral N. nipponensis Sakai, 1971, from Taiwan and Japan. spinules. Merus dorsoventrally compressed; extensor It differs from N. nipponensis chiefly in being less margin lined with numerous, short, subequal spines prominently spinose: the secondary spines covering in addition to paired distal spines; dorsal surface with the surfaces of the walking legs are distinctly shorter, irregular row of 4–7 major spines of similar size to and the antennal peduncle is only sparsely granulate distal extensor spines; flexor margin lined with short, or spinulate, rather than prominently spinulose. subequal spines in addition to 2–5 larger spines; major Neolithodes flindersi differs from N. brodiei in patterns spines on surfaces of merus distinctly standing out of spination, particularly of the ventral surfaces of the from secondary spinules. Carpus with 4 or 5 strong coxae of the walking legs, spination of the walking spines in addition to numerous small spines. Propodus leg dactyli, spination of the pereopod 2–4 meri and in dorsoventrally compressed; as long as or longer than morphometric features. These differences between N. merus; with 4–7 spines on extensor margin and 3–5 flindersi and N. brodiei are discussed under the account similar spines on dorsal surface; flexor margin with of the latter. numerous smaller spines. Dactylus curved, rounded As in other lithodids, dorsal spination of N. flindersi in cross section, with numerous small spines covering is most pronounced in smallest specimens, becoming proximal half (pereopod 2) to three-quarters (pere- progressively shorter with increasing size. Similarly, opods 3–4); apex corneous. the proportional length of the walking leg dactyli de- Pereopod 2 length 2.20–2.47 pcl (male), 1.84–2.24 creases with increasing size, ranging from about 0.8 pcl (female). Merus 0.74–0.81 pcl (male), 0.60–0.73 propodus length in the smallest specimen (juvenile pcl (female); length:height ratio 4.42–4.59 (male), male, pcl 21.6 mm) to 0.6 in the largest specimens (male, 3.21–3.98 (female). Carpus 0.54–0.55 merus length pcl 154.1 mm). The only known ovigerous female is (male), 0.56–0.60 (female). Propodus 0.93–0.99 merus 151.1 mm pcl, although females appear to be mature length (male), 0.95–0.99 (female); length:height ratio by about 95 mm pcl based on development of the abdo- 9.20–10.63 (male), 7.08–8.93 (female). Dactylus 0.62 men and presence of setae on the coxal surfaces. propodus length (male), 0.61–0.73 (female). One specimen of N. flindersi (AM P35606) was col- Pereopod 3 length 2.85–2.89 pcl (male), 2.26–2.74 lected from Chrysogorgia orientalis, and another (SAM pcl (female). Merus 0.84–0.94 pcl (male), 0.71–0.85 pcl C6857) from an unidentified black coral. The male (female); length:height ratio 4.56–4.91 (male), 3.50–4.09 holotype (NMNZ Cr11763) has an egg mass on the (female). Carpus 0.53–0.54 merus length (male), inside of the carapace in the left branchial chamber (egg 0.56–0.59 (female). Propodus 0.96–1.01 merus length diameter 3.98–4.08 mm). These eggs probably belong (male), 1.01–1.02 (female); length:height ratio 9.94– to a species of liparid fish (Careproctus sp.), which uses 10.65 (male), 7.94–10.02 (female). Dactylus 0.58–0.59 the crab as a mobile ‘home’ and uses the branchial propodus length (male), 0.62–0.68 (female). chambers of various lithodids for egg incubation.

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Plate 1. A, Lithodes aotearoa sp. nov., male holotype, pcl 88.6 mm, off Hawkes Bay (NIWA 34924). B, Lithodes aotearoa sp. nov., ovigerous female, pcl 118.9 mm, Chatham Rise (NIWA 42893). C, Lithodes australiensis sp. nov., male, pcl 60.3 mm, SSE of Southeast Cape (TM G3659). D, Lithodes rachelae sp. nov., male paratype, pcl 117.3 mm, Great Australian Bight (SAM C6367). E, Lithodes richeri Macpherson, 1991, ovigerous female, pcl 125.1 mm, SE of Brush Island (AM P35605). F, Lithodes robertsoni sp. nov., male holotype, pcl 128.1 mm, Chatham Rise (NIWA 42889). (Photo credits: A, P. Shearer, NIWA; C, K. Gowlett-Holmes, CSIRO; E, K. Lowe, Australian Museum; F, P. Marriott, NIWA).

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Plate 2. A, Neolithodes brodiei Dawson & Yaldwyn, 1970, ovigerous female, pcl 120.9 mm, Chatham Rise (NIWA 34923). B, Neolithodes bronwynae sp. nov., male holotype, pcl 123.3 mm, Whakatane Seamount (NIWA 49026). C, Neolithodes flindersi Ahyong, 2010, female, pcl 126.2 mm, E of Gabo Island (AM P52067). D, Neolithodes yaldwyni Ahyong & Dawson, 2006, male, pcl 116.9 mm, Ross Sea (NIWA 38209). E, Lithodes macquariae sp. nov. juvenile female, pcl 43.8 mm, Macquarie Ridge (NIWA 40902). F, Paralomis zealandica Dawson & Yaldwyn, 1971, female, pcl 79.4 mm, Chatham Rise (NIWA 42891). (Photo credits: C, K. Lowe, Australian Museum; D,E, P. Marriott, NIWA).

Plate 3 (opposite). A, Paralomis birsteini Macpherson, 1988a, male, pcl 69.5 mm, Ross Sea (NIWA 38505). B, Paralomis dawsoni Macpherson, 2001, male, pcl 120.8 mm, Challenger Plateau (NIWA 48671). C, Paralomis gowlettholmes sp.

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ nov., juvenile female paratype, pcl 15.9 mm, Tasmania (NMV J61054). D, Paralomis gowlettholmes sp. nov., ovigerous female, pcl 30.5 mm, Tasmania (TM G3607). E, Paralomis poorei sp. nov., juvenile male paratype, pcl 11.3 mm, Chatham Rise (NIWA 29526). F, Paralomis taylorae sp. nov., female holotype, pcl 39.8 mm, Tasmania (TM G3609). G, Paralomis stevensi Ahyong & Dawson, 2006, male, pcl 81.7 mm, Ross Sea (NIWA 27835). H, Paralomis staplesi sp. nov., male holotype, pcl 50.0 mm, Tasmania (NMV J61052). (Photo credits: A, E, P. Marriott, NIWA; B, S. O’Shea; C–D, F, H, K. Gowlett-Holmes, CSIRO).

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Plate 4. A, Lithodes chaddertoni sp. nov., holotype. B, Paralomis hirtella de Saint Laurent & Macpherson, 1998, Monowai Caldera, 25°48.237’S, 177°10.071’W, 1165 m. C, Lithodes aotearoa sp. nov., Kaikoura canyon, 42°35.01– 33.37’S, 173°43.81–43.77’E, 1379 m, TAN0616/95, 15 Nov 2006. D, Neolithodes yaldwyni Ahyong & Dawson, 2006, Balleny Islands, Ross Sea, 67°47.28–47.43’S, 179°45.71–47.33’W, 1072–1282 m, TAN0802/219, 4 Mar 2008. E, Par- alomis birsteini Macpherson, 1988a, Ross Sea, 68° 05.47–05.61’S, 179°16.63–14.96’W, 630–784 m, TAN0802/200, 2 Mar 2008. F, Paralomis zealandica Dawson & Yaldwyn, 1971, Chatham Rise, 44°00.37–00.31’S, 177°09.48–08.49’E, 637–630 m, TAN0705/54, 7 Apr 2007. (Photo credits: A, NMNZ; B, A. Rowden, NIWA; C–F, NIWA).

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Neolithodes flindersi occurs on soft substrates in outer Neolithodes yaldwyni Ahyong & Dawson, 2006 shelf and slope depths, and does not appear to range (Figs 59–62, Pl. 2D, 4D) beyond continental margins. It is frequently taken by deep-water trawlers in the central Great Austral- Neolithodes brodiei. — Thatje & Lörz, 2005: 335–336, fig. 2A ian Bight and is common between the Tasmanian [NIWA 3433, 3434 only; not N. brodiei Dawson & Yald- seamounts (Poore 2004; as N. brodiei). wyn, 1970]. Neolithodes yaldwyni Ahyong & Dawson, 2006: 46–54, figs 1–4 [type locality: N of Sturge Island, Balleny Islands, Distribution. Off Broken Bay, New South Wales, to Vic- 67°13.22–11.97’S, 164°17.78–14.87’E, 522–538 m]. toria, Tasmania and the Great Australian Bight, South — Frankham, 2009: 66. Australia; 887–1333 m, usually 950–1050 m.

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Type material. Holotype: NIWA 3434, male (cl.159 mm, tributed; merus ovate in cross-section, shorter than pcl 141.5 mm, cw 124.1 mm), N of Sturge Island, Balleny carapace; propodus subcylindrical in cross section; Islands, 67°13.22–11.97’S, 164°17.78–14.87’E, 522–538 dactylus distinctly longer than half propodus length. m, TAN0402/226, RV Tangaroa, 3 Mar 2004. Paratypes: NMNZ Cr11002, 2 males (cl 124.1–131.8 Description. Carapace: Pyriform, 1.02–1.15 times mm, pcl 101.7–103.6 mm, cw 91.6–95.2 mm), between Scott longer than wide; dorsal surface armed with short, and Balleny Islands, Ross Sea, 67°08–06’S, 170°54–47’E, stout spines and scattered, widely separated granules 900–1160 m, RV San Aotea II, trip 2010, haul 180, coll. E. or minute secondary spines; longest spine (hepatic Winslade, 14 Feb 2005; NIWA 3433, 1 male (cl 144.0 mm, spine) shorter than 0.2 pcl. Gastric region bearing 6 pcl 120.0 mm, cw 108.7 mm), seamount W of Sturge long spines forming transverse hexagon; with 2 near Island, Balleny Islands, 67°26.09–26.37’S, 163°52.98– transverse rows of 3 and 4 smaller spines, first across 51.79’E, 124–170 m, TAN0402/235, 4 Mar 2004; NMNZ centre of hexagon and second slightly anterior to Cr11003, 1 male (cl 183.3 mm, pcl 136.6 mm, cw 131.1 hexagon. Hepatic spine strong, anterodorsally directed; mm), near Scott Island, Ross Sea, 68°23–21’S, 179°53–57’E, margin between outer orbital spine and hepatic spine 1337–1050 m, RV San Aotea II, trip 2010, haul 137B, on with 2 or 3 short spines. Branchial regions with 8 or hook, B, coll. E. Winslade, 31 Jan 2005. 9 large dorsal spines in addition to scattered, smaller, secondary spines; margins with 13 large spines in ad- Other material examined. Ross Sea, Antarctica: NMNZ, dition to several shorter spines. Cardiac region with 4 2 males (cl +140–140.5 mm, pcl 101.7–112.7 mm, cw equal spines of similar size to gastric spines and pair of 99.1–97.6 mm), 65°S, 178°W, area 88.1C, FV San Aotea; shorter spines near intestinal region. Intestinal region NIWA 42890, 1 male (pcl 115.5 mm, cw 99.4 mm), Ross with 2 upright spines in transverse row and 2 smaller, Sea, area 88.1C, 65°07.20–08.40’S, 178°29.40–25.80’W, posterior directed spines. Pterygostomian region 1669–1950 m, trip 2530, set 11, coll. G. Higgins, 7 spinose or tuberculate, with small subdistal spine. Dec 2007; NMNZ Cr11780, 1 male (cl 136.4 mm, pcl Rostrum 0.12–0.25 pcl; median spine straight, nearly 109.00 mm, cw 95.3 mm), S of Scott Island, 68°02.00’S, horizontal, with pair of divergent dorsal spines and 179°44.00’W, 1312–1564 m, trip 1728, set 45, sample smaller pair of basal spines; ventral surface smooth. 65, FV Janas, coll. B. Fairhead & S. Smith, 21 Jan 2003; Posterior orbital near vertical in lateral view; outer NMNZ Cr11803, 1 ovigerous female (cl 135.3 mm, orbital spine slender, reaching to but not beyond cornea pcl 108.5 mm, cw 94.2 mm), SE of Balleny Islands, (when eyes directed forwards). Anterolateral spine 67°41.10’S, 167°08.50’E, 1307–1340 m, San Liberatore, about half to two-thirds length of outer orbital spine. Trip 1743, set 31, coll. G. Dolan, 11 Mar 2003; NMNZ Ocular peduncle: Longer than cornea; with scattered Cr11802, 1 male (pcl 109.3 mm, cw 99.8 mm), area 88.1, dorsal granules or spinules. 67°34.00–32.00’S, 167°18.00–13.00’E, 1290–1248 m, FV Antennule: Peduncle unarmed, reaching anteriorly San Liberatore, trip 1743, line 34, 12 Mar 2003; NMNZ beyond antennal peduncle by half to two-thirds length Cr11801, 1 male (cl 134.8 mm, pcl 120.5 mm, cw 113.0 of distal antennular peduncle article. mm), Ross Sea region, S of Antarctic Convergence, Antenna: Basal antennal article with small anterola- >70°S, Antarctic toothfish survey, 2000/2001; NIWA teral spine; outer margin of article 2 with small basal 38209, 1 male (pcl 116.9 mm, cw 106.1 mm), 68°06.52– spine and long slender spine that reaches beyond end 07.63’S, 179°14.30–15.36’W, 855–879 m, TAN0802/211, of article 4 but not to midlength of article 5; article 3 3 Mar 2008; NIWA 42888, 1 male (cl 170.4 mm, pcl with sharp to angular inner distal tooth; scaphocerite 146.1 mm, cw 135.6 mm), area 88.1K, 75°31.20’S, a minute, shorter than article 4, apex sharp to blunt; 170°10.80–27.00’W, 1177–1258 m, trip 2526/92, coll. article 4 unarmed, about half length of article 5. R. Coy, 5 Feb 2008. Abdomen: Somite 2 bearing 10 long spines and sev- Diagnosis. Carapace dorsal surface with thick, conical eral smaller spines on median plate; submedian plate spines and scattered, widely separated granules or with 4 or 5 long spines on posterior border and 2–4 minute secondary spines; longest spine not exceeding shorter spines on surface; lateral border of marginal 0.2 pcl. Rostrum 0.1–0.3 pcl; ventral surface smooth. plates with 6–8 teeth. Somites 3–5 multispinose or Posterior orbital margin near vertical; outer orbital nodulose in males; as calcified plates covered with spine slender, reaching to but not beyond cornea. stout, well-spaced spines in females. Telson spinose. Outer spine of antennal article 2 reaching beyond Egg diameter 2.70 mm. article 4; scaphocerite shorter than antennal article Pereopod 1 (chelipeds): Dimorphic but spination 4. Major cheliped palm with prominent dorsal and similar. Coxa with blunt tubercles and tufts of setae, lateral spines; dactylus with convex dorsal margin, unarmed; ischiobasis with 4 or 5 stout ventral spines. slightly longer than dorsal margin of palm. Walking Merus inner margin smooth or granular, with stout legs spinose, secondary spines absent or sparsely dis- subdistal spine; ventral margin with two rows of stout

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 59. Neolithodes yaldwyni Ahyong & Dawson, 2006, male, cl 159 mm, pcl 141.5 mm, cw 124.1 mm, N of Sturge Island, Balleny Islands (NIWA 3434). A, dorsal habitus. B, carapace. C, left antenna.

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 60 (above). Neolithodes yaldwyni Ahyong & Dawson, 2006, male, cl 159 mm, pcl 141.5 mm, cw 124.1 mm, N of Sturge Island, Balleny Islands (NIWA 3434). A, carapace, right lateral. B, ventral surface and abdomen. C, left pereopod 4 dactylus and carpus. D, posterior carapace and abdominal somite 2. E, right chela. F, left chela.

Figure 61 (opposite). Neolithodes yaldwyni Ahyong & Dawson, 2006, ovigerous female, pcl 108.5 mm, Ross Sea (NMNZ Cr11803). A, dorsal habitus. B, abdomen.

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ 105

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ spines, mesial row with 2 spines, lateral row with 4 Propodus 0.89–0.92 merus length (male), 0.98 (female); spines; dorsal and lateral surface spinose, spines largest length:height ratio 6.49–10.90 (male), 7.15 (female). distally. Carpus with prominent spines on dorsal and Dactylus 0.57–0.64 propodus length (male), 0.67 (fe- lateral surfaces; dorsal margin with row of 4 or 5 stout male). spines; lateral margin with 2 or 3 rows of 4 or 5 stout Pereopod 3 length 2.72–2.85 pcl (male), 2.06 pcl spines of similar size to dorsal row; ventral margin (female). Merus 0.94–1.01 pcl (male), 0.66 pcl (female); small, scattered, acute tubercles. Palm prominently length:height ratio 5.45–6.58 (male), 4.81 (female). spinose on dorsal, lateral and ventral surfaces, inner Carpus 0.49–0.52 merus length (male), 0.61 (female). surface with acute tubercles; dorsal margin with row Propodus 0.89–0.92 merus length (male), 0.90 (female); of about 5 prominent conical spines; midlateral surface length:height ratio 7.44–11.94 (male), 6.91 (female). with 2 rows of 4 or 5 spines of similar size to dorsal Dactylus 0.55–0.63 propodus length (male), 0.73 (fe- row; ventral surface with 2 rows of about 6 or 7 spines, male). smaller than lateral and dorsal spines. Pereopod 4 length 2.83–2.94 pcl (male), 2.13 pcl Major cheliped 1.32–1.55 pcl (male), 1.15 pcl (fe- (female). Merus 0.92–1.01 pcl (male), 0.67 pcl (female); male); upper palm length 1.13–1.29 times height (male), length:height ratio 5.68–7.12 (male), 5.06 (female). 1.19 (female); occlusal margins corneous for slightly Carpus 0.52–0.55 merus length (male), 0.61 (female). less than distal half, proximally with 3 calcareous Propodus 0.93–0.96 merus length (male), 7.64 (female); nodules; dactylus dorsal margin broadly convex, with length:height ratio 7.67–8.44 (male), 7.64 (female). Dac- rows of tufts of golden setae and proximal cluster of tylus 0.55–0.60 propodus length (male), 0.73 (female). 2 or 3 small spines, 1.15–1.26 times longer than dorsal margin of palm (male), 1.27 (female). Colour in life. Deep-red overall (Pl 2D). Minor cheliped 1.25–1.40 pcl (male), 1.15 pcl (female); upper palm length 1.42 times height (male), Remarks. Neolithodes yaldwyni Ahyong & Dawson, 2006, 1.30 (female); occlusal margins corneous for slightly from the Ross Sea, Antarctica, was recently described more than distal half, proximally crenulate; dactylus in detail based on male specimens. The first known dorsal margin broadly convex, with rows of tufts female of the species, a 108.5 mm pcl ovigerous speci- of golden setae and proximal cluster of 2 or 3 small men, is reported here. The female differs from males spines, 1.54–1.65 times longer than dorsal margin of in having stouter, thicker carapace spines, and as with palm (male), 1.57 (female). other lithodids, proportionally shorter pereopods. As Pereopods 2–4 (walking legs 1–3): Similar; segments in N. capensis, the female abdomen of N. yaldwyni is spinose, surface between major spines smooth or with composed of calcified plates covered with stout, well- few, minute, sparsely distributed spines, pereopod spaced spines. Notably, the dactyli of the walking legs 4 longest. Distal margins of coxae with short spines, are proportionally longer than in any of the males longest on coxa 3. Ischiobasis with 3 or 4 distal spines studied (0.7 versus 0.6 times propodus length). and 2 or 3 smaller ventral spines. Merus ovate in Neolithodes yaldwyni closely resembles N. capensis cross section, shorter than carapace; extensor margin Stebbing, 1905 (type locality: South Africa) in almost with 8 or 9 spines in addition to paired distal spines; all respects, but differs in having proportionally longer dorsal surface with row of 4–6 spines of similar size pereopod 2–4 dactyli. The pereopod 2–4 dactyli of N. to extensor spines; flexor margin with 2 rows of 5–7 capensis are half the propodus length in adult males spines. Carpus slightly longer than half merus length; versus 0.6 propodus length or greater in N. yaldwyni. extensor margin with 5 or 6 spines, distal and second Neolithodes capensis has been reported also from the Ker- proximal spines longest, about twice length of other guelen Islands (Macpherson 2004) and Bellingshausen spines; dorsal surface with 4 or 5 spines. Propodus Sea (García Raso et al. 2005). As remarked by Ahyong ovate to subcircular in cross section; with 7–9 spines & Dawson (2006), the Bellingshausen Sea record might on extensor margin and 8–10 similar spines on dorsal be referable to N. yaldwyni. At present, N. yaldwyni is surface; flexor margin with 6–10 smaller spines. Dac- known with certainty from localities in the Ross Sea tylus curved, rounded in cross section, with 4–6 small region, from the Scott and Balleny Islands to the Ross proximal spines, apex corneous; distinctly longer than Sea slope at 75°31’S. The affinities ofN . yaldwyni were half propodus length. discussed in detail by Ahyong & Dawson (2006). Pereopod 2 length 2.54–2.63 pcl (male), 1.91 pcl (female). Merus 0.85–0.92 pcl (male), 0.61 pcl (female); Distribution. Known only from the Ross Sea, Antarc- length:height ratio 5.15–6.27 (male), 4.19 (female). tica; 124–1950 m. Carpus 0.50–0.54 merus length (male), 0.64 (female).

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Paralomis White, 1856 P. cubensis Chace, 1939 [Caribbean Sea, western At- Paralomis White, 1856: 134 [type species: Lithodes granulosa lantic Ocean] Hombron & Jacquinot, 1846, pl. 8: figs 15–25. Gender: P. danida Takeda & Bussarawit, 2007 [Andaman Sea, feminine]. eastern Indian Ocean] Acantholithus Stimpson, 1858: 69 [type species: Lithodes histrix P. dawsoni Macpherson, 2001 [southwestern Pacific de Haan, 1844. Gender: masculine]. Ocean] Leptolithodes Benedict, 1895: 484 [type species: Paralomis P. debodeorum Feldmann, 1998 [extinct; Miocene, New aculeata Henderson, 1888, by subsequent designation Zealand] (Ahyong et al., 2010). Gender: masculine]. P diomedeae Pristopus Benedict, 1895: 486 [type species: Pristopus verrilli . (Faxon, 1893) [eastern Pacific Ocean] Benedict, 1895, by subsequent designation (Ahyong et al., P. dofleiniBalss, 1911 [western Pacific Ocean] 2010). Gender: masculine]. P. echidna sp. nov. [Australia, New Zealand] P. elongata Spiridonov, Türkay, Arntz & Thatje, 2006 Diagnosis. Carapace pentagonal or pyriform, not cov- [Bouvet Island, Southern Ocean] ering bases of walking legs; regions indicated; gastric P. erinacea Macpherson, 1988b [eastern Atlantic region elevated above other regions; cardiac region tri- Ocean] angular, separated from gastric region by deep groove; P. formosa Henderson, 1888 [southwestern Atlantic cervical groove shallow, indistinct. Rostrum comprised Ocean] of median spine and at least one pair of dorsal spines P. gowlettholmes sp. nov. [southwestern Australia] near base. Abdominal somite 2 comprising a single P. granulosa (Hombron & Jacquinot, 1846) [southwest- plate. Abdominal somites 3–5 comprising well-calci- ern Atlantic Ocean, southeastern Pacific Ocean] fied plates, without nodules and membranous areas; P. grossmani Macpherson, 1988b [eastern Atlantic each somite comprising median, paired submedian Ocean] and paired marginal plates (sometimes subdivided or P. haigae Eldredge, 1976 [southwestern Pacific Ocean] notched); marginal and submedian plates of somites P. hirtella de Saint Laurent & Macpherson, 1997 [south- 3 (and rarely 4) fused. Sternite 5 (between pereopods western Pacific Ocean] 2) without deep median fissure. Scaphocerite well- P. histrix (de Haan, 1844) [Japan, western Pacific developed, usually spinose. Walking legs (pereopods Ocean] 2–4) similar in form, walking leg 2 usually longest; P. hystrixoides Sakai, 1980 [Japan, western Pacific walking leg 3 always longer than pcl; dactyli of adults Ocean] with corneous spines along flexor margin. P. inca Haig, 1974 [southeastern Pacific Ocean] P. indica Alcock & Anderson, 1899 [northwestern In- Composition dian Ocean] P. aculeata Henderson, 1888 [Crozet and Prince Ed- P. investigatoris Alcock & Anderson, 1899 [northwestern wards Islands, southwest Indian Ocean, Southern Indian Ocean] Ocean] P. japonica Balss, 1911 [Japan, western Pacific Ocean] P. africana Macpherson, 1982 [Namibia, southeastern P. jamsteci Takeda & Hashimoto, 1990 [Japan, western Atlantic Ocean] Pacific Ocean] P. alcockiana Hall & Thatje, 2009b [South Carolina, P. kyushupalauensis Takeda, 1985 [Japan, western Pacific western Atlantic Ocean] Ocean] P. anamerae Macpherson, 1988b [Argentina, western P. longidactylus Birstein & Vinogradov, 1972 [Uruguay, Atlantic Ocean] southwestern Atlantic Ocean] P. arae Macpherson, 2001 [Fiji and Taiwan, western P. longipes Faxon, 1893 [Columbia to Peru, eastern Pacific Ocean] Pacific Ocean] P. arethusa Macpherson, 1994 [Barbados, western P. makarovi Hall & Thatje, 2009b [Bering Sea, northern Atlantic] Pacific Ocean] P. aspera Faxon, 1893 [central eastern Pacific Ocean] P. manningi Williams, Smith & Baco, 2000 [California, P. birsteini Macpherson, 1988a [Antarctic and Southern eastern Pacific Ocean] Ocean, Pacific sector] P. medipacifica Takeda, 1974 [western central Pacific P. bouvieri Hansen, 1908 [northeastern Atlantic Ocean] Ocean] P. mendagnai Macpherson, 2003 [Solomon Islands, P. ceres Macpherson, 1989 [western Indian Ocean] western Pacific Ocean] P. chilensis Andrade, 1980 [Chile, Peru, eastern Pacific P. microps Filhol, 1884 [eastern Atlantic Ocean] Ocean] P. multispina (Benedict, 1895) [northern Pacific P. cristata Takeda & Ohta, 1979 [Japan, western Pacific Ocean] Ocean] P. nivosa Hall & Thatje, 2009b [Philippines, western P. cristulata Macpherson, 1988b [eastern Atlantic Pacific Ocean] Ocean] 108

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ P. ochthodes Macpherson, 1988a [Indonesia, western range of carapace and pereopod forms, the genus is Pacific Ocean] united by the undivided plate of abdominal somite 2 P. odawarai (Sakai, 1980) [Japan, western Pacific and the calcified, rather than variously membranous, Ocean] plates of abdominal somites 3–5. Nevertheless, some P. otsuae Wilson, 1990 [Chile, southeastern Pacific potentially natural groups may be present within Ocean] Paralomis, such as the short-legged, densely spiny P. pacifica Sakai, 1978 [northwestern Pacific Ocean] forms similar to P. histrix and P. echidna, rela- P. papillata (Benedict, 1895) [eastern Pacific Ocean] tively less spiny, long-legged forms similar to P. pectinata Macpherson, 1988b [Venezuela, western P. aculeata and P. spinosissima, generally sparsely Atlantic Ocean] adorned forms similar to P. spectabilis and P. birsteini, P phrixa . Macpherson, 1992 [Peru, southeastern Pacific long-legged forms with cristate carapace margins Ocean] similar to P. verrilli and P. cristulata, and forms with P. poorei sp. nov. [New Zealand, Australia] a subhexagonal carapace densely covered by nodular P. roeleveldae Kensley, 1980 [South Africa, southwestern Indian Ocean] tubercles, such as P. granulosa, P. dawsoni, and P. dofleini. P. seagranti Eldredge, 1976 [southwestern Pacific Clearly, the range of forms within Paralomis is extensive Ocean] and phylogenetic analysis is much needed to further P. serrata Macpherson, 1988b [western Atlantic understand morphological diversity within the genus. Ocean] As discussed by Macpherson (1988b) and Ahyong et P. sonne Guzmán, 2009 [Chile, southeastern Pacific al. (2010), the taxonomic history of Paralomis has been Ocean] rather unstable, and three nominal genera are presently P. spectabilis Hansen, 1908 [northeast Atlantic Ocean] included in its synonymy: Acantholithus Stimpson, 1858 P. spinosissima Birstein & Vinogradov, 1972 [southwest (type species: Lithodes histrix de Haan, 1844), Lepto- Atlantic Ocean, Antarctica] lithodes Benedict, 1895, and Pristopus Benedict, 1895. P. staplesi sp. nov. [New Zealand, Australia] Benedict (1895) incorrectly regarded the type species P. stella Macpherson, 1988c [Reunion Islands, western of Paralomis, Lithodes granulosa Hombron & Jacquinot, Indian Ocean] 1846, as a species of Echinocerus White, 1848 (a junior P. stevensi Ahyong & Dawson, 2006 [Ross Sea, Ant- synonym of Lopholithodes Brandt, 1848), effectively arctica] sinking Paralomis. As such, Benedict (1895) erected two P. taylorae sp.nov. [southestern Australia] new genera to accommodate other species of Paralomis: P. truncatispinosa Takeda & Miyake, 1980 (= P. hetero- Leptolithodes and Pristopus. Leptolithodes was erected for tuberculata Tung, Wang & Li, 1984) [East China Sea, Paralomis aculeata Henderson, 1888, P. longipes Faxon, western Pacific Ocean] 1893, and two new species described therein, P. multi- P tuberipes . Macpherson, 1988a [Chile, southwestern spina Benedict, 1895 and P. papillata Benedict, 1895. Pacific Ocean] Pristopus was erected for two species, Pristopus verrilli P. verrilli (Benedict, 1895) [northern Pacific Ocean] Benedict, 1895, described therein as new, and Paralomis P. webberi sp. nov. [New Zealand, southwestern Pacific Ocean] formosa Henderson, 1888. Benedict (1895) did not des- P. zealandica Dawson & Yaldwyn, 1971 (= P. shinkai- ignate type species for either Leptolithodes or Pristopus, maruae Takeda & Hatanaka, 1984) [New Zealand, so in anticipation of further study of Paralomis, Ahyong southwestern Pacific Ocean] et al. (2010) selected Paralomis aculeata Henderson, 1888 and Pristopus verrilli Benedict, 1895 as the respective Remarks. Paralomis is the most speciose and most mor- type species for the two nominal genera. phologically diverse genus in the Lithodidae. Prior Paralomis includes the deepest living lithodid, to the present study, 62 extant and one fossil species P. bouvieri from the North Atlantic, which is known were recognised (De Grave et al. 2009; Hall et al. 2009). from as deep as 4152 m (Macpherson 1988b). Eleven species of Paralomis are known from the study area, of which six are new species: one is known only Key to adult Paralomis from New Zealand, from New Zealand (P. webberi sp. nov.), two exclusively Australia, and the Ross Sea from Australia (P. gowlettholmes sp. nov., P. taylorae sp. 1. Carapace densely covered with slender or conical nov.), and three occur in both regions (P. echidna sp. spines...... 2 nov., P. poorei sp. nov., P. staplesi sp. nov.). Paralomis – Carapace surface unarmed or with short, sparsely shinkaimaruae Takeda & Hatanaka, 1984 is shown to be a junior synonym of P. zealandica Dawson & Yaldwyn, distributed dorsal spines...... 6 1971. Thus, 67 extant species of Paralomis are currently 2. Meri of walking legs long, slender, pereopod 4 recognised. merus at least 0.85 pcl [southeastern Australia]..... The overall monophyly of Paralomis is not presently ...... P. taylorae sp. nov. in question (though further investigation is warranted into the the status of Lopholithodes with respect to – Meri of walking legs short, pereopod 4 merus less P. odawarai; Macpherson, 1988b) and despite the wide than 0.75 pcl...... 3

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ 3. Abdominal somites 4–6 with well-spaced granules 179°55’W, 1080 m]. — Dawson, 1989: 319. — Arana & and nodules [New Zealand and southeastern Aus- Retamal, 1999: 101 [Bellingshausen Sea records]. — Za- tralia]...... P. poorei sp. nov. klan, 2002: 770, 788. — Macpherson, 2004: 421, tab. 1. — Stevens, 2004: 6, 7 [part]. — Ahyong & Dawson, 2006: – Abdomen densely covered with slender spines or 55–58, figs 5–6, tab. 1. sharp conical tubercles in adults...... 4 Neolithodes brodiei. — Thatje & Lörz, 2005: 335 [NIWA 3432 4. Carapace covered with slender spines in juveniles only; not N. brodiei Dawson & Yaldwyn, 1970]. but low conical tubercles in adults [New Zealand] ...... P. zealandica Type material. Holotype: USNM 228830, female (cl 63.3, – Carapace covered with slender spines in adults.... pcl 55.0 mm, cw 53.1 mm), Ross Sea, N of Scott Island, ...... 5 67°29’S, 179°55’W, 1080 m, RV Eltanin, Stn 27-1946, 2 Feb 1967. 5. Subrostral lobe unarmed. Length of propodus of last walking leg about 3.7 times height in female Other material examined. Macquarie Ridge: NMV J61053, [Tasman Sea]...... P. echidna sp. nov. 1 male (pcl 64.8, cw 61.1 mm), N end of Gap, N of – Subrostral lobe multispinulate. Length of propodus Macquarie Island, 53°37.2’S, 159°11.3–17.4’E, with Bria- of last walking leg less than 4.4 times height in rosaccus callosus Boschma, 1158–1500 m, SS01/99/122, female [Kermadec Ridge]...... P. webberi sp. nov. 30 Jan 1999. 6. Carapace surface smooth, without spines or tuber- Ross Sea, Antarctica: NMNZ Cr11779, 1 male (cl cles; with tufts of stiff setae [hydrothermal vents; 89.9 mm, pcl 78.9 mm, cw 79.2 mm), N of Ross Sea, Fiji to Kermadec Ridge]...... P. hirtella 62°34.80’S, 173°35.40’E, 900 m, trip 1430, set 197B, FV – Carapace surface rugose, with spines or tubercles San Aotea II, 22 Apr 2001; NMNZ Cr11745–11749, 5 ...... 7 males (pcl 83.3–89.1 mm, cw 82.1–83.2 mm), Ross Sea, area 88.1C, 64°30.50’S, 171°13.8’W, 1128–1083 m, trip 7. Carapace surface entirely covered with blunt, wart- 1845, set 14, FV Janas, 21 Dec 2003; NIWA 23842, 1 male like tubercles; no dorsal spines [New Zealand to (cl 41.4 mm, cw 40.7 mm), 1 female (cl 35.2 mm, cw 32.5 Solomon Islands]...... P. dawsoni mm), both infected with Briarosaccus callosus Boschma, – Carapace surface with scattered short spines or 1930 (Rhizocephala), near Scott Island, 64°31.0–30.0’S, small tubercles; surface between spines and tuber- 171°14.0–08.0’W, 1225–1617 m, from stomach of Dis- cles generally smooth...... 8 sostichus mawsoni Norman, 1937, Avro Chieftain, set 8. Walking leg dactyli longer than dorsal extensor 24, 17 Dec 2003; NIWA 41869, 1 male (cl 86.0 mm, pcl margin of respective propodi...... 9 76.4 mm, cw 72.5 mm), Ross Sea, area 88.1C, 65°07.20– – Walking leg dactyli shorter than dorsal extensor 08.40’S, 178°29.40–25.80’W, 1669–1950 m, trip 2530, set margin of respective propodi...... 10 11, coll. G. Higgins, 7 Dec 2007; NIWA 3432, ovigerous female (cl 85.4 mm, pcl 73.9 mm, cw 72.5 mm), Bal- 9. Walking leg 3 propodus length:height ratio exceed- leny Seamount, 65°28.53–28.97’S, 161°02.88–02.75’E, ing 6.2 in males, exceeding 7.3 in females [Antarctica 760–750 m, TAN0402/269, RV Tangaroa, 7 Mar 2004; and Macquarie Ridge]...... P. birsteini NMNZ 11255, 1 male (cl 89.1 mm, pcl 78.2 mm, cw 73.8 – Walking leg 3 propodus length:height ratio 5.5 or mm), 66°35’S, 177°33’W, 1549 m, trip 1728, set 48, FV less in males, less than 5.6 in females [southeastern Janas; NIWA 38668, 1 male (cl 50.0 mm, pcl 42.6 mm, Australia]...... P. gowlettholmes sp. nov. cw 40.0 mm), 67°20.42–20.21’S, 179°55.91–56.53’W, 10. Carapace surface with scattered granules and few, 1130–1235 m, TAN0802/256, 8 Mar 2008; NIWA 24217, low conical spines. Cardiac region with 4 low tu- 1 male (cl 89.8 mm, cl 79.5 mm, cw 76.4 mm), near bercles [Ross Sea, Antarctica]...... P. stevensi Scott I., 67°21.01–21.02’S, 179°52.68–52.34’W, 540–600 – Carapace surface prominently multispinose. Car- m, sled tow, TAN0602/394, 6 Mar 2006; NIWA 38712, diac region with 4 spines [New Zealand and Aus- 1 ovigerous female (cl 69.6 mm, pcl 61.1 mm, cw 57.1 tralia]...... P. staplesi sp. nov. mm), 67°21.80–21.54’S, 179°57.11–57.25’W, 540–456 m, TAN0802/258, 8 Mar 2008; NIWA 38716, 1 male (cl 97.8 mm, pcl 90.9 mm, cw 86.6 mm), 1 ovigerous female (cl Paralomis birsteini Macpherson, 1988 69.9 mm, pcl 61.5 mm, cw 57.3 mm), 67°21.80–21.54’S, (Figs 63–66, Pl. 3A, 4E) 179°57.11–57.25’W, 540–456 m, TAN0802/258, 8 Mar 2008; NIWA 38505, 1 male (pcl 69.5 mm, cw 66.2 Paralomis spectabilis. — Birstein & Vinogradov, 1967: 390, figs mm), 67°22.92–22.67’S, 179°52.33–51.81’W, 420–456 1, 2. — Kirkwood, 1984: 37, fig. 45 [map only]. — Tkachuk, 1985: 1733–1734, figs A, B. — Zaklan, 2002: 773 [Ross Sea m, TAN0802/243, 7 Mar 2008; NMNZ Cr11001, 1 records] [not P. spectabilis Hansen, 1908]. male (cl 79.8, cw 74.7 mm), 1 female (cl 61.7, cw 59.4 Paralomis birsteini Macpherson, 1988a: 72–74, figs 4, 5a–e mm), both infected with Briarosaccus callosus Boschma, [type locality: Scott Island, Ross Sea, Antarctica, 67°29’S, 1930 (Rhizocephala), near Scott Island, 68°23–21’S,

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 63. Paralomis birsteini Macpherson, 1988a, male, pcl 79.4 mm, Ross Sea (NIWA 24217). A, dorsal habitus. B, carapace, right lateral view.

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 64. Paralomis birsteini Macpherson, 1988a, male, pcl 79.4 mm, Ross Sea (NIWA 24217). A, right chela. B, left chela. C, posterior carapace and abdominal somite 2. D, ventral surface and abdomen. E, anterior carapace, dorsal view. F, orbit and right antenna.

179°53–57’E, 1337–1050 m, RV San Aotea II, trip 2010, Diagnosis. Carapace subpentagonal, slightly wider than haul 137B, on hook, B.E. Winslade, 31 Jan 2005; NMNZ long; surface glabrous, sparsely covered with small, Cr11000, 1 male (cl 76.9, cw 73.4 mm), Ross Sea, Trip scattered granules and short conical spines, including 1862, haul 147, area 88.1G, RV San Aotea, obs. S. Voice median gastric spine; lateral margins with short conical [note: specific station data not available]. spines; outer distance between bases of anterolateral

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 65. Paralomis birsteini Macpherson, 1988a. A–B, male, pcl 79.4 mm, Ross Sea (NIWA 24217). C–D, ovigerous female, pcl 73.9 mm, Ross Sea (NIWA 3432). A, carapace. B, right pereopod 4 dactylus and propodus. C, dorsal habitus. D, abdomen.

spines slightly less than half carapace width. Rostrum times height of minor cheliped. Walking legs elongate, trispinose, broadest basally, without basal constriction. spinose, 2.2–2.7 pcl (male), 2.0–2.3 pcl (female); ventral Scaphocerite with 1 or 2 short inner basal spines and 1 spines of propodus widely spaced; dactyli longer than or 2 outer spines. Male chelipeds dimorphic, spinose; extensor margin of propodi. Walking leg 3 merus 0.7–0.8 major cheliped palm height less than 1.6 times that of pcl (male), 0.6–0.7 pcl (female), about 4–5 times longer minor cheliped. Major cheliped palm of female 1.2–1.3 than high; propodus length 6.2–7.9 times height.

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Description. Carapace: Subpentagonal, length 1.00–1.06 margins; dorsal and ventral margins spinose, inner times width; surface glabrous, sparsely covered with distal spine largest. Carpus with prominent spines on small, scattered granules. Gastric region convex, ele- dorsal and lateral surfaces; dorsal margin with row of vated, more prominent than other regions, with 5 short 3–5 spines; mesial margin with 2 spines, proximal larg- conical spines forming pentagon with anteriormost in est; ventral surface with 3 spines. Palm mesial margin midline. Lateral margin of hepatic region with 3 short with 6 spines, other surfaces with prominent, apically conical spines, anterior shorter. Branchial regions with setose tubercles or acute tubercles. Fingers with short 12–14 short marginal spines; surface with 4 or 5 low basal tubercle and rows of tufts of golden setae. conical spines in addition to low, scattered tubercles. Major cheliped 1.48–1.84 pcl (male), 1.38–1.44 Cardiac region subtriangular, with 2 pairs of short coni- (female); upper palm length 1.13–1.35 times height cal spines forming square. Intestinal region sparsely (male), 1.32–1.43 (female); occlusal margins of fingers granular. Pterygostomian region sparsely granular, corneous for distal third, proximally with 3 calcareous with prominent anterior spine. nodules; dactylus dorsal margin broadly convex, with Rostrum 0.13–0.17 pcl; broadest basally, without rows of tufts of golden setae and blunt proximal tooth, constriction; median spine smooth ventrally; paired 1.01–1.16 times longer than dorsal margin of palm dorsal spines divergent or subparallel, directed ob- (male), 1.33–1.34 (female). liquely upwards. Posterior orbital margin near con- Minor cheliped 1.44–1.66 pcl (male), 1.35–1.47 cave; outer orbital spine reaching to base of cornea. (female); upper palm length 1.18–1.38 times height Anterolateral spine as long as or shorter than outer (male), 1.35–1.50 (female); occlusal margin corneous orbital spine, with low marginal granule or small spine for slightly less than distal third, proximally crenulate; midway between outer orbital spine and anterolateral dactylus dorsal margin broadly convex, with rows of spine; outer distance between bases of anterolateral tufts of golden setae and blunt proximal tooth, 1.42– spines slightly less than half carapace width. 1.67 times longer than dorsal margin of palm (male), Ocular peduncle: Longer than cornea; with 2–5 dorsal 1.61–1.67 (female). granules. Pereopods 2–4 (walking legs 1–3): Similar, elongate, Antennule: Peduncle unarmed, reaching anteriorly spinose. Coxae smooth, unarmed; distal margins with beyond antennal peduncle by about three-quarters dense tufts of setae. Ischiobasis with 3–5 apically setose length of distal antennular peduncle article. ventral spines. Merus compressed, shorter than cara- Antenna: Basal antennal article with small ante- pace in both sexes; extensor margin with 8–11 spines in rolateral spine. Article 2 with angular to sharp inner addition to paired distal spines; dorsal surface with 4–8 distal margin; outer margin with small basal spine spines; flexor margin with rows of 5–7 spines; merus of and slender spine that does not reach beyond article pereopod 3 longer than that of pereopod 2 and pere- 4. Article 3 unarmed. Scaphocerite slender, reaching opod 4. Carpus slightly longer than half merus length, beyond midlength but not beyond distal three-quarters subcircular in cross section; extensor margin with 5 or of article 5, with 1 or 2 short inner spines and 1 or 2 6 spines; dorsal surface with 4–6 spines; flexor margin outer spines. Article 4 unarmed, about half length of unarmed. Propodus dorsoventrally flattened; slightly article 5. shorter than merus; with 9–14 spines on extensor mar- Abdomen: Ornamentation similar in both sexes. gin; dorsal surface with small scattered spines; flexor Somites sparsely granular. Somite 2 with small, widely margin with 5–8 spines. Dactylus broadly curved; separated spines along margins. Somite 3 median slightly longer than extensor margin of propodus; plate unarmed; submedian plates with low, irregular, surface with tufts of setae; extensor margin with 3 or triangular teeth on lateral margin; marginal plates 4 apically corneous spines proximally; lateral proximal absent, apparently undifferentiated. Somites 4–5 with surfaces with short, distinct sulcus, flanked ventrally unarmed median and submedian plates; marginal by 1 or 2 small, corneous spines; flexor margin lined plates undivided, with 2–4 low angular protrusions. with corneous spinules Somite 6 as long as or longer than wide, subquadrate, Pereopod 2 length 2.22–2.69 pcl (male), 2.01–2.25 with pair of small distal teeth; marginal plates short, pcl (female). Merus 0.72–0.85 pcl (male), 0.60–0.67 pcl triangular. Telson wider than long, apex rounded. (female); length:height ratio 4.05–5.04 (male), 4.01–4.67 Pereopod 1 (chelipeds): Spination similar in both sexes. (female). Carpus 0.55–0.58 merus length (male), 0.58– Major cheliped of males inflated, 1.39–1.57 times height 0.64 (female). Propodus 0.82–0.86 merus length (male), of minor cheliped; minor cheliped slender. Chelipeds 0.90–0.96 (female); length:height ratio 5.43–7.11 (male), of female unequal, major cheliped 1.24–1.25 times 6.69–6.95 (female). Dactylus 0.93–1.01 propodus length height of minor cheliped. Coxae smooth, unarmed; (male), 0.88–0.96 (female). distal margins with dense tufts of setae. Ischiobasis Pereopod 3 length 2.31–2.69 pcl (male), 2.10–2.33 with 4–8 stout, apically setose ventral spines. Merus pcl (female). Merus 0.73–0.86 pcl (male), 0.64–0.69 pcl with smooth mesial margins and tuberculate lateral (female); length:height ratio 4.08–5.26 (male), 4.81–4.99

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 66. Geographic distributions of Paralomis birsteini Macpherson, 1988 (○) and P. stevensi Ahyong & Dawson, 2006 (●).

(female). Carpus 0.55–0.57 merus length (male), 0.58– Colour in life. Light orange to orange-red; carapace 0.64 (female). Propodus 0.85–0.87 merus length (male), often paler than pereopods, especially on branchial 0.92–0.96 (female); length:height ratio 6.01–7.53 (male), regions (Pl. 3A). 6.81–7.66 (female). Dactylus 0.92–0.99 propodus length (male), 0.88–0.94 (female). Remarks. Paralomis birsteini Macpherson, 1988a has Pereopod 4 length 2.27–2.72 pcl (male), 2.09–2.34 been discussed extensively by Macpherson (1988a) and pcl (female). Merus 0.69–0.82 pcl (male), 0.60–0.68 pcl Ahyong & Dawson (2006). It closely resembles P. specta- (female); length:height ratio 4.26–5.23 (male), 5.01–5.11 bilis Hansen, 1908, P. stevensi Ahyong & Dawson, 2006, (female). Carpus 0.56–0.59 merus length (male), 0.58– and P. gowlettholmes sp. nov.; distinguishing features 0.64 (female). Propodus 0.91–0.94 merus length (male), are discussed under the accounts of P. gowlettholmes 0.97–1.02 (female); length:height ratio 6.22–7.84 (male), and P. stevensi (Table 3). 7.37–7.86 (female). Dactylus 0.90–0.96 propodus length The single specimen from Macquarie Ridge is a sig- (male), 0.89–0.94 (female). nificant northern range extension for the species. The Egg diameter 2.2–2.3 mm. specimen is parasitised by Briarosaccus callosus and, although a male, it has feminised pereopod morphology

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ in which the chelipeds are not significantly unequal, A preliminary molecular study found a paraphyletic and the walking leg morphometrics are in the range relationship between putative P. birsteini from the Bell- reported here for female P. birsteini from the Ross Sea. ingshausen Sea, Crozet and Kerguelen Islands (Thatje The smallest ovigerous female examined is 61.1 mm et al. 2008). Thus P. birsteini, as presently understood, pcl, and the smallest female overall (pcl 35.2 mm) has probably represents a species complex, and is currently an immature abdomen in which full asymmetry has under investigation in collaboration with S. Hall. The not yet been reached. The female holotype (pcl 55.0 present account of P. birsteini is based only on speci- mm) appears to be mature. mens collected at or near the type locality (i.e., Ross Paralomis birsteini has been attributed a near cir- Sea) and will serve as a basis for future comparison cum-Antarctic distribution, with records from the with P. birsteini from other localities. Several so-called Ross Sea in the vicinity of Scott and Balleny Islands, circum-Antarctic Crustacea, when more closely scru- the Crozet and Kerguelen Islands, the Bellingshausen tinised, have also proven to be composites, including Sea (Macpherson 1988b, 2004; Arana & Retamal 1999; various amphipods (Lörz et al. 2009), isopods (see Held García Raso et al. 2005; Ahyong & Dawson 2006), & Wägele 2005) and the king crab Lithodes murrayi (see and now Macquarie Ridge, near Macquarie Island. Macpherson 1988b, c; this study). Specimens from across the reported range of the species, however, have not been directly compared with Distribution. Known with certainty only from the Ross topotypic material. Notably, Arana & Retamal’s (1999) Sea, north to the Macquarie Ridge. Records from the figure of a specimen from the Bellingshausen Sea dif- Bellingshausen Sea, Crozet and Kerguelen Islands fers from P. birsteini sensu stricto, most significantly require confirmation; 420–1549 m in the Ross Sea, and in the walking leg dactyli being shorter rather than 620–1947 m outside of the Ross Sea (Macpherson 2004; longer than the respective propodal extensor margins. Thatje et al., 2008).

Table 3. Selected measurements of P. birsteini, P. gowlettholmes, P. stevensi, and P. webberi. Measurements in millimetres (mm). Abbreviations: postorbital carapace length (pcl).

Major cheliped Major cheliped Pereopod 2–4 Pereopod 4 Pereopod 4 Maximum Smallest height: minor height: minor dactylus length propodus propodus known ovigerous cheliped height cheliped height relative to length/ht length/ht pcl female (Male) (Female) extensor (Male) (Female) (pcl) propodal margin P. gowlettholmes .65–1.82 .35–1.46 Longer 4.96–5.53 3.67–5.59 50.1 28.3 P. birsteini .39–1.57 .24–1.26 Longer 6.22–7.84 7.37–7.86 90.9 61.1 P. stevensi .66–1.9 .31–1.33 Shorter 8.45–11.22 6.72–7.52 81.7 62.4 P. staplesi .59 ? 9.81 ? 50.0 ?

Paralomis dawsoni Macpherson, 2001 169°50.9’E, 820–1118 m, trip 1945, tow 27, 26 May (Figs 67–72A, 73, Pl. 3B) 2004. Challenger Plateau: NIWA 48671, 1 male (cl 140.6 mm, Paralomis sp. — Macpherson, 1990: 225, figs. 2c, 4. — O’Shea pcl 120.8 mm, cw 126.5 mm), 37°25.03’S, 168°06.06’E, et al., 1999: 49, fig. 16. — Zaklan, 2002: 810. 900 m, Z9724, coll. A. Knox, 3 Mar 1999. Paralomis sp. nov. — Zaklan, 2002: 773, 794. Bay of Plenty: NMNZ Cr11566–11569, 1 male (cl 127.9 Paralomis dawsoni Macpherson, 2001: 802–805, fig. 4 [type locality: New Caledonia]. — Webber & Naylor, 2004b: mm, pcl 114.7 mm, cw 117.8 mm), 2 ovigerous females 62. — Naylor et al., 2005: 46. — Ahyong et al., 2007: 156. (cl 116.7–122.1 mm, pcl 102.3–107.4 mm, cw 104.3–109.4 — Dawson, 2008: 7, fig. 8. mm), 1 female (cl 132.4 mm, pcl 114.7 mm, cw 121.1 Paralomis sp. [“multipustulata”]. — Dawson, 2008: fig. 6. mm), Bay of Plenty, FV Savannah, coll. B. Roughlin; NMNZ Cr11767, 1 female (pcl 142.7 mm, cw 139.8 mm), Type material. Holotype: MNHN Pg4278, ovigerous Bay of Plenty, coll. Gregg Gibbs. female (pcl 71.0 mm, cw 72.8 mm), New Caledonia, Hawkes Bay: NMNZ Cr5642, 1 male (cl 129.9 mm, seaward side of reef, from trap. pcl 112.0 mm, cw 118.4 mm), Ritchie Bank, 39°28.39’S, 178°23.57’E, 400–600 m, FV Betty T, coll. P. Terry, 30 Other material examined. West Norfolk Ridge: NMNZ Sep 1986. Cr12028, 1 ovigerous female (cl 128.1 mm, pcl 112.6 Louisville Ridge: NMNZ Cr11730, 1 male (cl 94.3+ mm, cw 114.7 mm, W of Ninety Mile Beach, 34°49.5’S, mm, pcl 85.4 mm, cw 89.7 mm), 40°58’S, 165°04’W, 800

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 67. Paralomis dawsoni Macpherson, 2001, male, pcl 120.8 mm, Challenger Plateau (NIWA 48671). A, dorsal habitus. B, carapace, right lateral view.

m, FV Peterson, coll. M. Miranovich, 2 Mar 1995. cold seep, 40°02.31–02.47’S, 178°08.58–08.62’E, 730–747 Hikurangi Margin: NIWA 29295, 1 ovigerous female m, TAN0616/06, 4 Nov 2006. (cl 105.8 mm, pcl 91.9 mm, cw 92.3 mm), Rock Garden Chatham Rise: NIWA 3905, 1 female (cl 80.6+ mm,

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 68. Paralomis dawsoni Macpherson, 2001, male, pcl 120.8 mm, Challenger Plateau (NIWA 48671). A, carapace. B, anterior carapace, dorsal view.

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 69. Paralomis dawsoni Macpherson, 2001, male, pcl 120.8 mm, Challenger Plateau (NIWA 48671). A, right chela. B, left chela. C, left pereopod 4. D, right antenna. E, ventral surface and abdomen. F, posterior carapace and abdominal somite 2. pcl 72.9 mm, cw 71.8 mm), top of Morgue Seamount, m, FV San Waitaki, Trip 1702/140, coll. T. Brunning & 42°43.17–43.00’E, 179°57.78–58.24’E, 986 m, baited pot, M. Leslie, 16 Oct 2002; NIWA 54363, 1 juvenile male (cl TAN0104/193, Z10715, 19 Apr 2001; NMNZ Cr11969, 12.6 mm, pcl 10.3 mm, cw 9.0 mm), 44°01.67–01.87’S, 1 ovigerous female (cl 118.5 mm, pcl 104.9 mm, cw 174°35.46–35.44’E, 801–823 m, TAN0905/121, 28 Jun 108.1 mm), 42°48.2–48.4’S, 179°48.7–49.1’W, 866–970 2009.

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 70. Paralomis dawsoni Macpherson, 2001, ovigerous female, pcl 91.9 mm, Rock Garden cold seep (NIWA 29295). A, dorsal habitus. B, abdomen.

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Diagnosis. Carapace subhexagonal, about as long as long, slender, slightly overreaching distal peduncular wide; surface without spines, uniformly covered with article; with multiple lateral spines of varying length, nodulose, minutely setose, wart-like, microscopically occasionally secondarily bifurcate; dorsomesially with setose, compound tubercles, tubercles smallest anterior 2 or 3 spines, shorter than half length of lateral spines. to level of hepatic spines; margins microscopically se- Article 4 unarmed, about half length of article 5. tose; hepatic to anterior branchial margins lined with Abdomen: Ornamentation similar in both sexes. short but prominent conical spines; lateral branchial Somites 2–3 covered with nodulose tubercles. Somites margins subparallel, straight to slightly concave; 4–6 covered with short conical spines; margins dentate posterior branchial margins nodular or with short to spinose. Telson spinose; semicircular. acute tubercles or spines. Rostrum trispinose, dorsal Pereopod 1 (chelipeds): Strongly spinose and tuber- pair of spines prominent, without basal constriction. culate, setose, unequal; ornamentation similar on both Scaphocerite multispinose. Abdomen covered with sides and in both sexes. Coxal margins tuberculate. wart-like, compound tubercles. Chelipeds and walk- Ischiobasis tuberculate laterally and ventrally. Merus ing legs covered with compound tubercles and coarse outer surfaces granular; mesial surface relatively spines. smooth, with strong inner distal spine, flanked by several small spines; dorsal surface granular proximally Description. Carapace: Subhexagonal, 0.95–1.02 times as becoming spinose distally. Carpus surfaces tuberculate; long as wide; regions distinct, gastric region more in- dorsal margin with 6–8 strong, upright spines; ventral flated than others; surface uniformly covered with nod- margin with short, conical spines and acute tubercles. ulose, minutely setose, wart-like compound tubercles, Palm surfaces tuberculate, dorsal margin spinose; inner tubercles smallest anterior to level of hepatic spines. (mesial) surface sparsely tuberculate, apically setose. Hepatic and anterior branchial margins divergent (ap- Major cheliped 1.69–1.93 pcl (male), 1.20–1.23 proximately to level of pereopod 3); with 9–15 conical, (female); upper palm length 0.99–1.05 times height microscopically setose spines of varying length. Lat- (male), 1.01–1.08 (female); occlusal margins of fingers eral branchial margins subparallel, straight or slightly corneous for distal third, proximally with 3 calcareous convave, granular. Posterior branchial margin arcuate, nodules; dactylus dorsal margin broadly convex, with tuberculate, usually unarmed, occasionally with acute rows of tufts of golden setae and blunt proximal tooth, tubercle anteriorly (largest male also with 2–4 short 1.04–1.12 times longer than dorsal margin of palm acute tubercles along margin). Pterygostomian region (male), 1.27–1.31 (female). with small anterior spine; surface with small, coarse Minor cheliped 1.85–1.61 pcl, (male), 1.17–1.21 granules, densest and largest on posterior half. (female); upper palm length 1.00–1.06 times height Rostrum 0.11–0.16 pcl; broad basally, not constricted (male), 1.03–1.11 (female); occlusal margin corneous proximal to dorsal spines; median spine curved slightly in distal third to half, proximally crenulate to weakly dorsally, overreaching outerorbital spine; with blunt dentate; dactylus dorsal margin broadly convex, with ventral tooth; paired dorsal spines short, divergent, rows of tufts of golden setae and blunt proximal tooth, directed obliquely upwards; with single small upright 1.59–1.64 times longer than dorsal margin of palm spine at base of paired dorsal spines. Posterior orbital (male), 1.85–1.91 (female). margin concave, unarmed; outer orbital spine slender, Pereopods 2–4 (walking legs 1–3): Similar, elongate, directed anteriorly, overreaching eyes, with or with- spinose. Pereopod 2 longest. Coxae distal margins out small inner basal spine. Anterolateral spine half irregular; ventral surfaces tuberculate, becoming low as long as outer orbital spine; outer distance between and blunt in mature females and obsolete on pereopod bases of anterolateral spines distinctly less than half 3–4. Ischiobasis spinose. Merus and carpus dorsal sur- carapace width. faces tuberculate (also spinose on pereopod 4), exten- Ocular peduncle: Longer than cornea; dorsally with sor and flexor margins prominently spinose; ventral short, scattered granules or spines, distalmost spine surfaces unarmed, at most with a row of low tubercles longest; distal spine about half corneal diameter, or small spines near flexor and extensor margins. Pro- slightly overreaching cornea. podus shorter than merus; tuberculate and spinose on Antennule: Peduncle unarmed, reaching anteriorly all surfaces, spines longest on dorsal margins. Dactylus beyond apex of antennal peduncle by two-thirds length curved, laterally compressed; longer than carpus; with of distal antennular peduncle article. cluster of proximal spines and scattered tufts of golden Antenna: Basal antennal article with acute granules setae; flexor margin lined with 9–13 obliquely inclined and slender, arcuate anterolateral spine. Article 2 corneous spinules; apex corneous. with slender inner spine and short dorsal spine; outer Pereopod 2 length 1.87–1.95 pcl (male), 1.34–1.46 margin with granules and 2 small spines proximally, pcl (female). Merus 0.59–0.66 pcl (male), 0.40–0.45 pcl and long, distal spine overreaching article 3. Article 3 (female); length:height ratio 3.45–3.71 (male), 2.56–2.96 unarmed. Scaphocerite multispinose; primary spine (female). Carpus 0.61–0.63 merus length (male), 0.68–

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 71. Paralomis dawsoni Macpherson, 2001. A, female, pcl 142.7 mm, Bay of Plenty (NMNZ Cr11767). B, male, pcl 112.0, Ritchie Bank (NMNZ Cr5642).

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 72. A, Paralomis dawsoni Macpherson, 2001, juvenile, pcl 10.3 mm, Chatham Rise (NIWA 54363). B–E, Paralomis seagranti Eldredge, 1976, male, cl 63.5 mm, pcl 58.1 mm, cw 63.9 mm, South Bank, Guam (ZRC 2000.0738). A–B, dorsal habitus. C, right pereopod 4. D, right anterior branchial margin of carapace. E, carapace, right lateral view.

0.77 (female). Propodus 0.75–0.76 merus length (male), 3.86–4.24 (female). Dactylus 0.91–1.03 propodus length 0.78–0.79 (female); length:height ratio 4.23–4.39 (male), (male), 0.94–1.00 (female). 3.24–3.76 (female). Dactylus 0.93–1.01 propodus length (male), 0.98–1.05 (female). Colour in life. Uniform orange-red to red-brown (Pl. Pereopod 3 length 1.81–1.94 pcl (male), 1.33–1.46 3B). pcl (female). Merus 0.56–0.62 pcl (male), 0.41–0.45 pcl (female); length:height ratio 3.37–3.45 (male), 2.54–2.75 Remarks. Paralomis dawsoni was first formally recorded (female). Carpus 0.64–0.65 merus length (male), from New Zealand by Webber & Naylor (2004b). It is 0.69–0.75 (female). Propodus 0.81 merus length (male), the largest species of the genus in New Zealand waters 0.82–0.83 (female); length:height ratio 4.54–4.90 (male), and the 142.7 mm pcl female from the Bay of Plenty is 3.65–4.04 (female). Dactylus 0.90–1.02 propodus length the largest known specimen of the species. Allometric (male), 0.92–1.04 (female). variation within the material at hand of P. dawsoni is mi- Pereopod 4 length 1.70–1.88 pcl (male), 1.30–1.31 nor in comparison to ‘spiny’ species of Paralomis such pcl (female). Merus 0.49–0.56 pcl (male), 0.36–0.36 pcl as P. zealandica and P. histrix. The primary changes are (female); length:height ratio 3.15–3.58 (male), 2.73–3.01 in the size and sharpness of the spines along the antero- (female). Carpus 0.68–0.71 merus length (male), 0.72– lateral carapace margins (proportionally shortest in the 0.81 (female). Propodus 0.90–0.91 merus length (male), largest specimens) and in the degree of concavity of the 0.91–0.92 (female); length:height ratio 4.74–5.20 (male), branchial margin at the level of pereopods 3 and 4, be-

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ ing least concave in the largest specimens. The smallest broadly hexagonal carapace outline with spines along specimen, a juvenile male (pcl 10.3 mm), differs from the hepatic and at least the anterior branchial margin, adults chiefly in the following respects: the marginal and in which the carapace and abdomen are densely carapace spines are proportionally longer; the posterior covered with blunt, rounded, compound tubercles, branchial margin bears several prominent spines; the such as P. alcockiana Hall & Thatje, 2009b [South Caro- compound tubercles of the carapace and abdomen are lina, western Atlantic], P. cubensis Chace, 1939 [Carib- less developed and do not fully crowd the surface; and bean Sea], and P. seagranti Eldredge, 1976 [Guam, west- several short conical spines are present on the carapace ern Pacific].Paralomis dawsoni is readily distinguished surface (three gastric including median; two cardiac; from P. alcockiana by the absence of the prominently two branchial). The dorsal carapace spines of the early enlarged median gastric tubercle on the carapace, and juvenile are completely lost in adults; and the posterior from P. cubensis by having much more prominent cara- branchial spines are lost or reduced to acute tubercles pace tubercles with margins well-demarcated from the in some adult males, lost in adult females. adjacent carapace surface. In P. cubensis, the carapace As in other species of Paralomis, sexual dimorphism tubercles are relatively low with ill-defined margins. is evident in the more strongly inflated chelipeds and Of the aforementioned species, P. dawsoni most closely proportionally longer walking legs in adult males. Ad- resembles P. seagranti (Fig. 72B–E) from Guam and Ki- ditionally, the fingers of the minor cheliped of the males ribati in bearing spines along the hepatic and anterior above 110 mm pcl exhibit a distinct gape, rather than branchial margins, with margins unarmed posterior occluding for the full length as in smaller specimens. to the level of the gastro-cardiac groove. Paralomis Paralomis dawsoni resembles those species with a dawsoni differs from P. seagranti in more pronounced

Figure 73. Geographic distribution of Paralomis dawsoni Macpherson, 2001.

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ and nodular carapace tubercles (rather than relatively base of scaphocerite. Scaphocerite a long, slender flat), a lower, less strongly inflated gastric region, in the spine overreaching cornea spines; with lateral and concave rather than evenly convex midlateral branchial dorsal spines. Abdominal somites densely covered margins, and in lacking the long, dense, golden setae with slender spines, becoming shorter on successive covering the marginal carapace spines. The two species somites. Chelipeds short, stout, with outer and upper also differ in the degree of setation of the pereopods: surfaces densely spinose; mesial surface of palm with sparse in P. dawsoni and dense in P. seagranti, giving low, scattered, setose tubercles, otherwise smooth; the latter a distinctly hirsute appearance. dactyli proximally spinose. Walking legs similar, short, Paralomis dawsoni and P. zealandica are the two most stout, not exceeding 1.8 pcl (male), 1.4 pcl (female); common species of the genus in New Zealand. Their segments densely spinose except for ventral surface ranges are largely discrete, with P. dawsoni a northern of meri and coxae; pereopod 4 merus length less than form, and P. zealandica a southern form, with their 3.0 times height (male), 2.5 (female); propodus length ranges overlapping at the Subtropical Convergence in about 4.3 times height (male), 3.7 (female); dactylus the vicinity of the Chatham Rise. shorter than carpus, surface spinose for proximal half. The ovigerous female of P. dawsoni from Rock Gar- Somite 6 length 1.08–1.15 times width. den cold seep (NIWA 29295) is the first lithodid to be recorded from a New Zealand cold seep. Description. Carapace: Pyriform, 1.06–1.13 times longer than wide; regions distinct; surface and margins Distribution. New Caledonia, the Solomon Islands and densely and uniformly covered with slender spines northern New Zealand; 400–1118 m. The New Zealand of similar length; surface of spines with scattered, range includes the West Norfolk Ridge and Challenger minute, well-spaced simple setae; longest spine (on Plateau in the west, and in the east, the Bay of Plenty, anterior branchial margin) 0.27 pcl; cardiac region Ritchie Bank, and the Louisville Ridge, south to the with 27 (male) or 18–22 (female) spines; cervical groove Chatham Rise. distinct. Pterygostomian region with slender anterior spine; posterior two-thirds with field of spinules; Paralomis echidna sp. nov. (Figs 74–81) surface of anterior third smooth except for scattered granules near dorsal margin and with cluster of 6 or 7 Paralomis cf. histrix. — Poore, 2004: 269, fig. 76e. blunt spinules anteriorly. Rostrum 0.24–0.25 pcl; broad basally, not constrict- Type material. Holotype: NMNZ Cr9285, ovigerous fe- ed proximal to dorsal spines; median spine slender, male (cl 41.9 mm, pcl 33.6 mm, cw 29.6 mm), Gascoyne ventral lobe triangular with blunt apex, unarmed; Seamount, Tasman Sea, 36°30’S, 156°30’E, about 650 m, dorsally with 1 or 2 median spines and pair of laterally in baited pots, coll. J. Cave, Feb 1992. divergent spines directed obliquely upwards. Posterior Paratype: NMV J15122, ovigerous female (cl + 47.4 orbital margin concave, unarmed; outer orbital spine mm, pcl 38.8 mm, cw 33.5 mm), 75 km SE of Point slender, directed anterolaterally, reaching midlength Hicks, Victoria, 38°11.50’S, 149°56.50’E, 636 m, Engels of cornea. Anterolateral spine as long as outer orbital high-lift demersal trawl, RV Soela, S06/84, 28 Nov spine; outer distance between bases of anterolateral 1984. spines slightly less than half carapace width. Ocular peduncle: Longer than cornea; with scattered Other material examined. Norfolk Ridge: NMNZ granules and 9–12 dorsal spines, anterior 3 or 4 the Cr16862, 1 ovigerous female (pcl 32.1 mm, cw 28.5 longest, anteriorly directed, arranged in arcuate row mm), Norfolk Ridge, S of Wanganella Bank, 33°41.0’S, above cornea; longest spine overreaching cornea by 167°52.6’E, 817 m, trawl, FV Seamount Explorer, coll. J. more than length of cornea. Houston, 13 Mar 2006. Antennule: Peduncle unarmed, reaching anteriorly No specific locality: NMNZ Cr11636, 1 male (cl 61.7 beyond antennal peduncle by length of distal anten- mm, pcl 49.7 mm, cw 46.9 mm), no data, coll. G. McIn- nular peduncle article. ninnie. Antenna: Basal antennal article with slender, curved, anterolateral spine, not reaching base of scaphocerite. Diagnosis. Carapace pyriform, slightly longer than Article 2 with inner distal spine, sometimes bearing 1 wide; surface and margins densely and uniformly or 2 subdistal granules; outer margin with 3–5 spines, covered with slender spines of similar length; outer distalmost longest, reaching midlength of ultimate distance between bases of anterolateral spines slightly peduncular article. Article 3 unarmed. Scaphocerite a less than half carapace width. Rostrum ventral lobe long, slender spine overreaching cornea spines; lateral rounded, unarmed. Ocular peduncle dorsally spinu- margins with 2 or 3 long spines; dorsally with 1–4 small late, spinules overreaching cornea. Basal antennal spines; inner margin unarmed. Article 4 unarmed, article with slender anterolateral spine reaching to almost half length of article 5.

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 74. Paralomis echidna sp. nov., ovigerous female holotype, cl 41.9 mm, pcl 33.6 mm, cw 29.6 mm, Gascoyne Seamount (NMNZ Cr9285). A, dorsal habitus. B, ventral habitus.

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 75. Paralomis echidna sp. nov., ovigerous female holotype, cl 41.9 mm, pcl 33.6 mm, cw 29.6 mm, Gascoyne Seamount (NMNZ Cr9285). A, right chela, outer surface. B, right chela, inner surface. C, left chela, outer surface. D, carapace, right lateral view. E, right pereopod 4, dorsal view. F, right pereopod 4, ventral view.

Abdomen: Ornamentation similar in both sexes. less than half-length of dorsal carapace spines. Somite Somites densely covered with slender spines, becom- 6 length 1.08–1.15 times width. Telson subtriangular ing shorter on successive somites. Spines on somite 2 with rounded apex, with 5–7 blunt, apically setose similar to dorsal carapace spines. Spines of somites 3–6 tubercles.

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 76. Paralomis echidna sp. nov., ovigerous female holotype, cl 41.9 mm, pcl 33.6 mm, cw 29.6 mm, Gascoyne Seamount (NMNZ Cr9285). A, anterior carapace, dorsal view. B, anterior carapace, right lateral view. C, carapace, anterior view. D, posterior carapace and abdomen. E, abdomen, posterior somites and telson. F, abdomen.

Pereopod 1 (chelipeds): Densely spinose, unequal. spinose laterally, ventrally with clusters of golden Major cheliped 1.65 times height of minor cheliped setae. Merus with smooth mesial surface, other sur- (male), 1.48–1.56 (female). Coxae unarmed, setose; faces spinose. Carpal surfaces densely spinose except distal margins with dense tufts of setae. Ischiobasis for mesial surface with clusters of golden setae. Palm

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 77. Paralomis echidna sp. nov., ovigerous female holotype, cl 41.9 mm, pcl 33.6 mm, cw 29.6 mm, Gascoyne Seamount (NMNZ Cr9285). A, right antenna, ventral view. B, right antenna, dorsal view. C, rostrum, right lateral view. D, right eye, lateral view. E, right pereopod 4 dactylus. Scale = 1.0 mm.

surfaces densely spinose except for mesial surface with row of 2–5 small setose tubercles near flexor mar- with low scattered tubercles bearing tufts of golden gin. Carpus spinose, longest spines as high as carpus. setae. Fingers with tufts of golden setae; dactyli with Propodus dorsoventrally flattened; all surfaces spinose, proximal cluster of 2–4 spines. longest spines twice propodal height. Dactylus curved; Major cheliped 1.73 pcl (male), 1.21–1.35 (female); shorter than carpus; surface spinose for proximal half; upper palm length 1.02 times height (male), 0.94–1.08 distal half with tufts of golden setae; flexor margin (female); occlusal margins corneous for distal quarter, lined with 7–9 obliquely inclined corneous spinules; proximally with 4 low calcareous prominences; dacty- apex corneous. lus dorsal margin broadly convex, with rows of tufts Pereopod 2 length 1.79 pcl (male), 1.33–1.38 pcl of golden setae and proximal cluster of 2–4 spines, (female). Merus 0.58 pcl (male), 0.41–0.43 pcl (female); 1.07 times longer than dorsal margin of palm (male), length:height ratio 2.88 (male), 2.45–2.51 (female). 1.30–1.34 (female). Carpus 0.71 merus length (male), 0.76–0.78 (female). Minor cheliped 1.64 pcl (male), 1.21–1.26 (female); Propodus 0.85 merus length (male), 0.86–0.91 (female); upper palm length 1.06 times height (male), 1.13–1.18 length:height ratio 4.60 (male), 3.84–3.96 (female). Dac- (female); occlusal margins corneous for distal third tylus 0.74 propodus length (male), 0.75–0.83 (female). to half, proximally crenulate; dactylus dorsal margin Pereopod 3 length 1.74 pcl (male), 1.30–1.34 pcl broadly convex, with rows of tufts of golden setae and (female). Merus 0.58 pcl (male), 0.39–0.40 pcl (female); proximal cluster of 2–4 spines, 1.53 times longer than length:height ratio 2.95 (male), 2.36–2.39 (female). dorsal margin of palm (male), 1.58–1.67 (female). Carpus 0.72 merus length (male), 0.78–0.84 (female). Pereopods 2–4 (walking legs 1–3): Similar, short, stout, Propodus 0.85 merus length (male), 0.91–0.98 (female); densely spinose; pereopod 2 longest. Coxae, unarmed, length:height ratio 4.64 (male), 3.52–3.83 (female). Dac- surface setose, distal margins crenulate. Ischiobasis tylus 0.72 propodus length (male), 0.74–0.84 (female). spinose. Merus ovate in cross section, extensor and Pereopod 4 length 1.56 pcl (male), 1.20–1.28 pcl flexor surfaces spinose, longest extensor spines at about (female). Merus 0.48 pcl (male), 0.35–0.37 pcl (female); distal third; ventral surface smooth, unarmed, at most length:height ratio 2.91 (male), 2.43–2.52 (female).

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 78. Paralomis echidna sp. nov., male, cl 61.7 mm, pcl 49.7 mm, cw 46.9 mm, no locality (NMNZ Cr11636). A, dorsal habitus. B, ventral habitus.

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 79. Paralomis echidna sp. nov., male, cl 61.7 mm, pcl 49.7 mm, cw 46.9 mm, no locality (NMNZ Cr11636). A, right chela. B, left chela. C, anterior carapace, left lateral view. D, left pereopod 4. E, carapace, right lateral view. F, posterior carapace and abdominal somite 2.

Carpus 0.80 merus length (male), 0.84–0.88 (female). Egg diameter: 2.0–2.3 mm at ‘black eye-spot’ Propodus 0.91 merus length (male), 1.01–1.07 (female); stage. length:height ratio 4.34 (male), 3.68–3.73 (female). Dac- tylus 0.74 propodus length (male), 0.74–0.81 (female). Colour in life. Not known.

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 80. Paralomis echidna sp. nov., ovigerous female, pcl 32.1 mm, cw 28.5 mm, Norfolk Ridge (NMNZ Cr16862). A, dorsal habitus. B, ventral habitus.

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Etymology. Name echidna, the common name of stouter in the male. The male further differs from the the spiny Australian and Papua New Guinean females in bearing 30 instead of 18–22 cardiac spines. monotremes; used as a noun in apposition. Paralomis echidna and P. webberi also appear to mature at a smaller size than P. histrix. Both P. echidna and P. Remarks. Paralomis echidna sp. nov. most closely resem- webberi are ovigerous at less than 35 mm pcl, whereas bles P. histrix (de Haan, 1844) from Japan, and P. web- a Japanese specimen of P. histrix examined here (ZLKU beri sp. nov. from the Kermadec Ridge, which together 13247; Fig. 82) appears to have just reached maturity are readily distinguished from other congeners by the at 68.6 mm pcl. densely spinose carapace, abdomen, chelipeds and Unfortunately, the specific provenance of the only walking legs, and the relatively short walking legs in known male of P. echidna, collected by a New Zealand which the meri do not exceed 0.6 pcl in males or 0.5 commercial fisherman based in Tauranga, is not known pcl in females. Paralomis echidna differs from P. histrix (R. Webber, pers. comm.), though it was probably in the pattern of carapace and abdominal spination in collected from northern New Zealand waters. The which most of the adjacent spines are of approximately female from the Norfolk Ridge and the male differ equal length, and in having a comparatively wider from southern Tasman Sea specimens of P. echidna in frontal region, with the distance between the bases of bearing a single spine in the midline of the rostrum, the anterolateral spines only slightly rather than dis- rather than two, and in having proportionally shorter tinctly less than half carapace width. In contrast, the dorsal carapace spines. The shorter carapace spines carapace and abdominal ornamentation of P. histrix in the single known male might be attributable to its is composed of numerous long spines interspersed larger body size, but the Norfolk Ridge female is of with numerous short, apically rounded, anteriorly similar size to the southern Tasman Sea specimens. recurved spines of approximately half the length of The Norfolk Ridge female also differs from southern the longer spines (Fig. 81; see also Ikeda 1998: pl. 73). Tasman Sea specimens in having 22 instead of 18 or Also, the carapace spines of P. echidna (and P. webberi) 19 cardiac spines. The aforementioned differences are are minutely and sparsely setose rather than glabrous presently regarded as intraspecific variation, although as in P. histrix. Features distinguishing P. echidna from further study might reveal these differences to be sig- P. webberi are outlined under the account of the latter. nificant. For this reason, the type series is restricted to Differences between the male and female P. echidna are the southern Tasman Sea specimens. chiefly sexually dimorphic and allometric: the walk- Poore (2004) reported P. echidna as P. cf. histrix. The ing legs are more elongate and the right cheliped is only species presently known from Australia that P.

Figure 81. Geographic distributions of Paralomis echidna sp. nov. (▲), P. hirtella de Saint Laurent & Macpherson, 1997 (○), P. poorei sp. nov. (□), P. staplesi sp. nov. (■), and P. webberi sp. nov. (●).

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 82. Paralomis histrix (de Haan, 1844), female, cl 85.0 mm, pcl 68.6 mm, cw 66.7 mm, Kochi Prefecture, Japan (ZLKU 13247). A, carapace. B, anterior carapace, dorsal view. C, dorsal habitus. E, right posterior carapace surface.

echidna is likely to be confused with is P. poorei (also Distribution. Presently known only from the Tasman Sea known from New Zealand), a species also densely cov- between Victoria, Australia, the Gascoyne Seamount, ered with spines. Paralomis echidna can be immediately and the southern Norfolk Ridge; 636–817 m. distinguished from P. poorei by its spinose rather than tuberculate abdominal somites 4–6.

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Paralomis gowlettholmes sp. nov. 38.4 mm, cw 37.9 mm), 1 ovigerous female (cl 39.2 mm, (Figs 83–87, Pl. 3C–D) pcl 34.5 mm, cw 33.3 mm), approximately 46.5 nautical miles [86 km] SSE of South East Cape, 44°22.7’S, Paralomis sp. MoV2716. — Poore et al., 1998: 66, 72. 147°07.3’E, 1050–1170 m, trawled, coll. K. Gowlett-Hol- Paralomis cf. birsteini. — Poore, 2004: 269, fig. 76b. mes, 12 Feb 1992; NMV J44020, 1 juvenile male (cl 11.5 mm, pcl 9.91 mm, cw 8.21 mm), D1 seamount, 97.4 km Type material. (All Tasmania). Holotype: SAM C6859, SSE of South East Cape, 44°24.0’S, 147°19.2’E, 1900 m, male (pcl 50.1 mm, cw 49.1 mm), ovigerous female (cl RV Southern Surveyor, SS01/97 stn 22, epibenthic sled, 52.5 mm, pcl 45.42 mm, cw 43.64 mm), approx. 425 nau- coll. T.N. Stranks et al., 25 Jan 1997. tical miles [787 km] SE of South East Cape, 44°09.2’S, 147°31.8’E, 970–1120 m, trawled, FV Belinda, coll. K. Diagnosis. Carapace subpentagonal, slightly wider than Gowlett-Holmes, 1 Feb 1992. long; surface glabrous, sparsely covered with small, Paratypes: SAM C6860, ovigerous female (cl 52.5 scattered granules and short conical spines, including mm, pcl 45.4 mm, cw 43.6 mm), approx. 425 nauti- median gastric spine; lateral margins with short conical cal miles [787 km] SE of South East Cape, 44°09.2’S, spines; outer distance between bases of anterolateral 147°31.8’E, 970–1120 m, trawled, FV Belinda, coll. K. spines slightly exceeding half carapace width. Rostrum Gowlett-Holmes, 1 Feb 1992; NMV J44018, 2 oviger- trispinose, broadest basally, without basal constriction. ous females (cl 34.1–45.3 mm, pcl 28.78–39.0 mm, cw Scaphocerite with 1 or 2 short inner basal spines and 1 28.13–37.42 mm), J1 seamount, 82.6 km SSE of South or 2 outer spines. Male chelipeds dimorphic, spinose; East Cape, 44°14.4’S, 147°21.6’E, 1200 m, RV Southern major cheliped palm height 1.7–1.8 times that of mi- Surveyor, SS01/97 stn 40, epibenthic sled, coll. T.N. nor cheliped. Major cheliped palm of female 1.4–1.5 Stranks et al., 27 Jan 1997; SAM C6861, 1 male (cl 44.9 times height of minor cheliped. Walking legs elongate, mm, pcl 40.7 mm, cw 37.7 mm), 1 female (cl 41.5 mm, spinose, length 2.0–2.6 pcl; ventral spines of propodus pcl 34.5 mm, cw 33.6 mm), approximately 46 nauti- widely spaced; dactyli longer than extensor margin cal miles [85 km] SE of South East Cape, 44°14.8’S, of propodi. Walking leg 3 merus 0.7–0.8 pcl (male), 147°27.5’E, 1081–1130 m, trawled, coll. K. Gowlett-Hol- 0.5–0.6 pcl (female), less than 5 times longer than high; mes, 12 Feb 1992; SAM C6862, 1 female (cl 46.3 mm, pcl propodus length 3.7–5.6 times height. 39.9 mm, cw 38.0 mm), approximately 44 nautical miles [81 km] SSE of South East Cape, 44°15.2’S, 147°21.2’E, Description. Carapace: Subpentagonal, 1.01–1.09 times 1050–1120 m, trawled, FV Belinda, coll. K. Gowlett- longer than wider; surface glabrous, sparsely covered Holmes, 12 Feb 1992; NMV J44013, 1 juvenile female (cl with small, scattered granules. Gastric region convex, 10.2 mm, pcl 8.5 mm, cw 7.8 mm), 1 crushed specimen elevated, more prominent than other regions, with 5 (cl approx. 19 mm), Sister 1 seamount, 82.9 km SSE short conical spines forming pentagon with anterior- of South East Cape, 44°16.2’S, 147°17.4’E, 1100 m, RV most in midline. Lateral margin of hepatic region with 3 Southern Surveyor, SS01/97 stn 15, epibenthic sled, coll. short conical spines, anterior shorter. Branchial regions T.N. Stranks et al., 23 Jan 1997; TM G3583, 1 female (cl with 8 or 9 short marginal spines; surface with 4 low 28.9 mm, pcl 23.2 mm, cw 22.1 mm), Hill K1, approx. conical spines in addition to low, scattered tubercles. 88 km SSE of Southeast Cape, 44°17’S, 147°25’E, 1600 Cardiac region subtriangular, with 2 pairs of short coni- m, epibenthic sled, RV Southern Surveyor, SS0197/17, cal spines forming square. Intestinal region sparsely 23 Jan 1997; NMV J44016, 1 female (cl 33.5 mm, pcl granular. Pterygostomian region sparsely granular, 28.3 mm, cw 26.8 mm), Dory Hill seamount, 83.2 km with prominent anterior spine. SSE of South East Cape, 44°19.2’S, 147°07.2’E, 1280 m, Rostrum 0.1–0.2 pcl; broadest basally, without con- RV Southern Surveyor, SS01/97 stn 49, epibenthic sled, striction; median spine smooth ventrally, occasionally coll. T.N. Stranks et al., 29 Jan 1997; NMV J61054, 1 ju- with small proximal dorsal spine or tubercle; paired venile female (cl 19.5 mm, pcl 15.9 mm, cw 14.5 mm), dorsal spines divergent, directed obliquely upwards. 44°19.78’S, 146°53.23’E, 1305 m, FV Tommy Thompson, Posterior orbital margin near concave; outer orbital ROV Jason 2, TN228-J2-382-016-004, 17 Dec 2008; TM spine reaching to base of cornea. Anterolateral spine G3607, 1 ovigerous female (cl 35.8 mm, pcl 30.5 mm, as long as outer orbital spine, often with low marginal cw 28.6 mm), hill U, approximately 85 km SSE of South granule midway between outer orbital spine and ante- East Cape, 44°20’S, 147°10’E, 1083 m, RV Southern Sur- rolateral spine; outer distance between bases of antero- veyor, SS0197/34, CSIRO, 27 Jan 1997; NMV J44019, 1 lateral spines slightly exceeding half carapace width. ovigerous female (cl 48.7 mm, pcl 41.4 mm, cw 40.27 Ocular peduncle: Longer than cornea; with 2–5 dorsal mm), U seamount, 86.4 km SSE of South East Cape, granules. 44°20.4’S, 147°09.6’E, 1250 m, RV Southern Surveyor, Antennule: Peduncle unarmed, reaching anteriorly SS01/97 stn 35, epibenthic sled, coll. T.N. Stranks et beyond antennal peduncle by about three-quarters al., 27 Jan 1997; SAM C6863, 1 male (cl 49.0 mm, pcl length of distal antennular peduncle article.

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 83. Paralomis gowlettholmes sp. nov., male holotype, pcl 50.1 mm, cw 49.1 mm, SE of South East Cape, Tasmania (SAM). A, dorsal habitus. B, carapace. E, right pereopod 4.

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 84. Paralomis gowlettholmes sp. nov., male holotype, pcl 50.1 mm, cw 49.1 mm, SE of South East Cape, Tas- mania (SAM). A, right chela. B, left chela. C, ventral surface and abdomen. D, posterior carapace and abdominal somite 2. E, carapace, right lateral view. F, anterior carapace, dorsal view. Antenna: Basal antennal article with small ante- and 1–3 outer spines. Article 4 unarmed, about half rolateral spine. Article 2 with angular to sharp inner length of article 5. distal margin; outer margin with small basal spine and Abdomen: Ornamentation similar in both sexes. slender spine not reaching beyond article 4. Article Somites sparsely granular. Somite 2 with small, widely 3 unarmed. Scaphocerite slender, reaching beyond separated spines along margins. Somite 3 median plate midlength of article 5, with 1 or 2 short inner spines unarmed; submedian plates with low, irregular, trian-

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 85. Paralomis gowlettholmes sp. nov., female paratype, cl 33.5 mm, pcl 28.3 mm, cw 26.8 mm, Dory Hill seamount, Tasmania (NMV J44016). A, dorsal habitus. B, ventral surface and abdomen.

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 86. Paralomis gowlettholmes sp. nov. A, juvenile female paratype (cl 19.5 mm, pcl 15.9 mm, cw 14.5 mm (NMV J61054). B, juvenile female paratype (pcl 23.2, cw 22.1 mm), Hill K1 (TM G3583).

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ gular teeth on lateral margin; marginal plates absent, with 3 or 4 apically corneous spines proximally; lateral apparently undifferentiated. Somites 4–5 with un- proximal surfaces with short, distinct sulcus, flanked armed median and submedian plates; marginal plates ventrally by 1 or 2 small, corneous spines; flexor margin undivided, with irregular triangular teeth. Somite 6 lined with corneous spinules slightly longer than wide, subquadrate, with pair of Pereopod 2 length 2.12–2.62 pcl (male), 1.78–1.96 small distal teeth; marginal plates short, triangular. pcl (female). Merus 0.70–0.82 pcl (male), 0.58–0.63 pcl Telson wider than long, apex rounded. (female); length:height ratio 3.84–4.48 (male), 3.42–4.47 Pereopod 1 (chelipeds): Spination similar in both (female). Carpus 0.55–0.59 merus length (male), 0.61– sexes. Major cheliped of males strongly inflated, 0.66 (female). Propodus 0.80–0.84 merus length (male), 1.7–1.8 times height of minor cheliped; minor cheliped 0.86–0.93 (female); length:height ratio 4.29–5.62 (male), slender. Chelipeds of female unequal, major cheliped 3.98–5.81 (female). Dactylus 0.94–1.00 propodus length 1.4–1.5 times height of minor cheliped. Coxae smooth, (male), 0.85–0.99 (female). unarmed; distal margins with dense tufts of setae. Is- Pereopod 3 length 2.07–2.64 pcl (male), 1.87–1.99 chiobasis with 4–8 stout, apically setose ventral spines. pcl (female). Merus 0.72–0.81 pcl (male), 0.59–0.65 pcl Merus with smooth mesial margins and tuberculate lat- (female); length:height ratio 3.96–4.74 (male), 3.41–4.50 eral margins; dorsal and ventral margins spinose, inner (female). Carpus 0.54–0.59 merus length (male), 0.57– distal spine largest. Carpus with prominent spines on 0.65 (female). Propodus 0.82–0.89 merus length (male), dorsal and lateral surfaces; dorsal margin with row of 0.89–0.99 (female); length:height ratio 4.86–5.87 (male), 3–5 spines; mesial margin with 2 spines, proximal larg- 4.28–6.06 (female). Dactylus 0.92–0.94 propodus length est; ventral surface with 3 spines. Palm mesial margin (male), 0.83–0.96 (female). with 6 spines, other surfaces with prominent, apically Pereopod 4 length 2.02–2.38 pcl (male), 1.81–2.08 setose tubercles or acute tubercles. Fingers with short pcl (female). Merus 0.66–0.79 pcl (male), 0.49–0.60 pcl basal tubercle and rows of tufts of golden setae. (female); length:height ratio 3.73–4.38 (male), 2.69–4.76 Major cheliped 1.77–1.96 pcl (male), 1.30–1.55 (female). Carpus 0.57–0.61 merus length (male), 0.63– (female); upper palm length 1.11–1.14 times height 0.68 (female). Propodus 0.89–0.91 merus length (male), (male), 1.23–1.40 (female); occlusal margins of fingers 0.96–1.03 (female); length:height ratio 4.96–5.53 (male), corneous for distal third, proximally with 3 calcareous 3.67–5.59 (female). Dactylus 0.93–0.95 propodus length nodules; dactylus dorsal margin broadly convex, with (male), 0.81–0.96 (female). rows of tufts of golden setae and blunt proximal tooth, 1.09–1.17 times longer than dorsal margin of palm Colour in life. Pinkish-red (Pl. 3C–D). (male), 1.15–1.25 (female). Minor cheliped 1.47–1.75 pcl (male), 1.16–1.59 Etymology. Named for Karen Gowlett-Holmes, CSIRO (female); upper palm length 1.22–1.30 times height Marine Research, who collected the holotype of the (male), 1.38–1.54 (female); occlusal margin corneous new species, in addition to many other specimens for slightly less than distal third, proximally crenulate; studied herein. dactylus dorsal margin broadly convex, with rows of tufts of golden setae and blunt proximal tooth, 1.49– Remarks. Paralomis gowlettholmes sp. nov. closely re- 1.62 times longer than dorsal margin of palm (male), sembles P. spectabilis Hansen, 1908 (type locality: off 1.42–1.68 (female). Iceland), P. birsteini Macpherson, 1988a, and P. stevensi Pereopods 2–4 (walking legs 1–3): Similar, elongate, Ahyong & Dawson, 2006 (both with type locality: Ross spinose. Coxae smooth, unarmed; distal margins with Sea) in carapace ornamentation in which the primary dense tufts of setae. Ischiobasis with 3–5 apically setose spines are low and simple with the surface between ventral spines. Merus compressed, shorter than cara- the primary spines smooth or sparsely granulate. The pace in both sexes; extensor margin with 8–11 spines in new species is readily distinguished from P. spectabilis addition to paired distal spines; dorsal surface with 4–8 and P. stevensi by the length of the dactyli of the walk- spines; flexor margin with rows of 5–7 spines; merus ing legs—slightly longer, rather than shorter than the of pereopod 3 slightly longer than that of pereopod 2 extensor margin of the propodi in P. gowlettholmes. and pereopod 4. Carpus slightly longer than half merus Paralomis gowlettholmes further differs from P. stevensi length, subcircular in cross section; extensor margin in the having much shorter walking legs in males, and with 5 or 6 spines; dorsal surface with 4–6 spines; further differs from P. spectabilis in the armed inner flexor margin unarmed. Propodus dorsoventrally flat- margin of the scaphocerite. tened; slightly shorter than merus; with 9–14 spines Paralomis gowlettholmes is most similar to P. birst- on extensor margin; dorsal surface with small scat- eini in the proportional lengths of the walking leg tered spines; flexor margin with 5–8 spines. Dactylus dactyli, but differs chiefly in the proportional size of broadly curved; slightly longer than extensor margin of the major chela and the less slender walking legs. As propodus; surface with tufts of setae; extensor margin in P. stevensi, the right chela of male P. gowlettholmes is

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considerably larger than the left, with the right palm stevensi, both of which are known to exceed 80 mm pcl height about 1.7–1.8 versus less than 1.6 times or less compared to 50.1 mm pcl for the largest specimen of that of the left in P. birsteini. The right chela of females the new species. The smallest ovigerous female of P. is also comparatively larger in P. gowlettholmes, being gowlettholmes is 28.3 mm pcl, less than half the size of about 1.4–1.5 times higher than the left versus 1.1–1.2 the smallest ovigerous P. birsteini (pcl 61.1 mm) and P. in P. birsteini. The walking legs of P. gowlettholmes are stevensi (pcl 62.4 mm). Moreover, the abdomen of the noticeably stouter than those of P. birsteini, quantified smallest female P. birsteini examined (pcl 35.2 mm), in the length:height ratio of the propodi. In P. gowlet- although parasitised, has not acquired full asymmetry tholmes, the length:height ratio of the walking leg 3 suggesting that females are immature at that size. propodus does not exceed 5.6 but is greater than 6.2 in The similar morphology and Southern Ocean P. birsteini (see Table 3). The difference in the length: distribution of Paralomis birsteini, P. gowlettholmes and height ratio between P. gowlettholmes and P. birsteini P. stevensi points to a close relationship between the is even more pronounced between females, <5.6 and three species. >7.3 respectively. Paralomis gowlettholmes appears to mature earlier Distribution. Presently known only from southern and attain a smaller size than either P. birsteini or P. Tasmania; 970–1900 m.

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Paralomis hirtella de Saint Laurent & 1 male (cl 62.9 mm, pcl 58.3 mm, cw 64.6 mm; with Macpherson, 1997 rhizocephalan Briarosaccus callosus), 25°48.29–48.59’S, (Figs 81, 88–91, Pl. 4B) 177°10.19–09.82’E, 1140–1054 m, TAN0411/06, 3 Oct 2004; NIWA 32233, 1 male (cl 50.8 mm, pcl 46.6 mm, Paralomis hirtella de Saint Laurent & Macpherson, 1997: 722– cw 49.6 mm; with rhizocephalan Briarosaccus callosus), 2 726, figs 1–4 [type locality: Lau Basin, 22°13’S, 176°38’W, juveniles (cl 7.7–7.9 mm), 25°48.33’S, 177°10.18’W, 1102 1750 m]. — Macpherson & Segonzac in Desbruyères et al. m, SO192-2/18, 3 May 2007; NIWA 32224, 4 males (cl 2006: 436. — Dawson, 2008: 4–8, figs 1–5. 28.9–46.7 mm, pcl 22.3–42.7 mm, cw 23.9–45.8 mm), Paralomis aff. jamsteci. — Webber & Naylor, 2004b: 63. 5 females (cl 22.0–46.0 mm, pcl 19.9–41.0 mm, cw 20.7–42.7 mm), 25°48.33’S, 177°10.18’W, 1041 m, RV Type material. Holotype: MNHN Pg4658, male (pcl 34.9 Sonne, SO192-2/20, 3 May 2007; NIWA 32223, 1 male mm, cw 32.0 mm), Lau Basin, Valufa Rise, Hine Hina (cl 34.5 mm, pcl 31.5 mm, cw 33.1 mm), 25°48.21’S, site, 22°13’S, 176°38’W, 1750 m, 14–18 May 1989. 177°10.08’W, 1197 m, RV Sonne, SO192-2/21, 3 May 2007. Other material examined. New Zealand, Brothers Seamount: NIWA 18518, 1 juvenile male (cl 10.2 mm, pcl Diagnosis. Carapace subpentagonal, as long as or slight- 8.7 mm, cw 8.8 mm), 1 juvenile female (cl 16.2 mm, pcl ly wider than long; lateral margins with 12–14 short, 14.3 mm, cw 15.2 mm), Brothers Seamount, 34°51.64’S, slender, well-spaced anterolaterally directed spines; 179°03.47’E, 1572 m, KOK0507/21, 14 May 2005; NIWA surface devoid of spines, smooth, unarmed, bearing 34926, 1 male (cl 62.7 mm, pcl 58.4 mm, cw 61.0 mm), numerous tufts of golden setae. Rostrum broad basally, 34°51.66’S, 179°03.42’E, 1603 m, DPH0501/72, crab without constriction, extending to or slightly beyond pot, CRB 11; NIWA 4078, 1 male (cl 36.9 mm, pcl 32.8 cornea; ventrally unarmed; paired dorsal spines short, mm, cw 34.8 mm), 1 indeterminate juvenile (cl 9.7 mm, directed anteriorly, occasionally reduced to low setose pcl 8.3 mm, cw 8.4 mm), 34°51.756’S, 179°03.476’E, nodules. Scaphocerite with small inner and 1 or 2 outer 1649 m, DSV Shinkai 6500, 6K#852, RK6-B-71, 27 spines. Abdominal somites unarmed, with scattered Oct 2004; NIWA 4079, 1 male (cl 54.5 mm, pcl 50.2 tufts of setae. Cheliped palms unarmed except for low mm, cw 51.3 mm), 34°52.723’S, 179°04.304’E, 1336 m, teeth on dorsal margin. Walking legs unarmed except 6K#854, RK6-B-71, 1 Nov 2004; NIWA 34925, 7 males for row of blunt spines on meral extensor margins. (cl 40.7–57.6 mm, pcl 36.5–51.8 mm, cw 38.3–56.0 mm), 8 ovigerous females (cl 41.1–57.9 mm, pcl 36.8–52.6 Description. Carapace: Subpentagonal, about as long as mm, cw 39.6–57.0 mm), 34°52.9’S, 179°04.13’E, 1208 wide, length 0.90–1.04 width; regions distinct, gastric m, DPH0501/104; NIWA 34932, 1 male (cl 59.2 mm, region more inflated than others; surface smooth, pcl 53.4 mm, cw 55.1 mm), 1 ovigerous female (cl 37.9 unarmed, bearing numerous tufts of golden setae. mm, pcl 34.3 mm, cw 35.9 mm), 34°52.9’S, 179°04.13’E, Anterior margin almost transverse, usually with small 1208 m, DPH0501/104; NIWA 3918, 2 males (cl spine between outerorbital and anterolateral spine 18.1–29.3 mm, pcl 15.7–25.8 mm, cw 16.2–27.4 mm), and occasionally with spine at inner base of anterola- 34°52.96–52.69’S, 179°04.02–04.93’E, 1197–1538 m, teral spine. Lateral margins with 12–14 short, slender, TAN0107/141, 22 May2001; NIWA 3914, 2 females, well-spaced anterolaterally directed spines. Posterior larger ovigerous (cl 30.5–38.8 mm, pcl 27.2–35.4 mm, margin relatively straight, slightly convex to slightly cw 28.9–38.2 mm), 34°53.12–53.35’S, 179°04.49–05.93’E, concave. Pterygostomian region with small anterior 1197–1526 m, TAN0107/136, 21 May 2001. spine; surface irregular, unarmed. Monowai Caldera: NIWA 18002, 1 male (cl 47.7 mm, Rostrum short, 0.07–0.20 pcl; broad basally, not pcl 42.8 mm, cw 45.3 mm; with rhizocephalan Bria- constricted proximal to dorsal spines; median spine rosaccus callosus), 1 ovigerous female (cl 33.6 mm, pcl horizontal or ventrally deflexed, overreaching outer- 30.7 mm, cw 32.0 mm), 25°48.24’S, 177°10.07’W, 1168 orbital spine, extending to or slightly beyond cornea; m, KOK0505/8, 8 Apr 2005; NIWA 18003, 1 juvenile ventrally unarmed; paired dorsal spines short, directed male (cl 8.1 mm, pcl 6.7 mm, cw 6.5 mm), 25°48.33’S, anteriorly, occasionally reduced to low setose nodules. 177°10.25’W, 1050 m, KOK0505/3, 7 Apr 2005; NIWA Posterior orbital margin concave, unarmed; outer or- 18004, 1 male (cl 66.5 mm, pcl 61.1 mm, cw 66.8 mm; bital spine slender, directed anteriorly, not overreach- with rhizocephalan Briarosaccus callosus), 25°48.25’S, ing eyes. Anterolateral spine subequal to outer orbital 177°10.11’E, 1143 m, KOK0505/14, 10 Apr 2005; NIWA spine; outer distance between bases of anterolateral 18009, 3 males (cl 16.6–41.6 mm, pcl 14.8–36.9 mm, cw spines slightly exceeding half carapace width. 30.7–36.9 mm), 2 females (cl 33.7–53.2 mm, pcl 30.8–49.3 Ocular peduncle: Longer than cornea; dorsally with mm, cw 33.0–51.2 mm; largest with rhizocephalan Bria- tufts of golden setae, unarmed. rosaccus callosus), 25°48.29–48.59’S, 177°10.19–09.82’E, Antennule: Peduncle unarmed, reaching anteriorly 1140–1054 m, TAN0411/06, 3 Oct 2004; NIWA 18010, beyond apex of antennal peduncle by half length of

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 88. Paralomis hirtella de Saint Laurent & Macpherson, 1997, male, pcl 53.4 mm, Brothers Seamount (NIWA 34932). A, dorsal habitus. B, anterior carapace, dorsal view.

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 89. Paralomis hirtella de Saint Laurent & Macpherson, 1997, male, pcl 53.4 mm, Brothers Seamount (NIWA 34932). A, right chela. B, left chela. C, right pereopod 4. D, right carapace, lateral view. D, ventral surface and abdomen. E, posterior carapace and abdominal somite 2.

distal antennular peduncle article. ing to or beyond midlength of article 5; with small inner Antenna: Basal antennal article unarmed. Article 2 denticle and 1 or 2 outer spines, distal longer. Article unarmed on inner and dorsal surface; outer margin 4 unarmed, about half length of article 5. with 1 or 2 acute granules proximally, and long, over- Abdomen: Ornamentation similar in both sexes. Sur- reaching end of article 3 but not overreaching article 4. face smooth, covered in tufts of golden setae; margins Article 3 with a few acute granules. Scaphocerite reach- smooth, unarmed. Telson spinose; semicircular.

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 90. Paralomis hirtella de Saint Laurent & Macpherson, 1997, female, pcl 34.3 mm, Brothers Seamount (NIWA 34932). A, dorsal habitus. B, ventral surface and abdomen.

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 91. Paralomis hirtella de Saint Laurent & Macpherson, 1997, Brothers Seamount (NIWA 18518). A, male juvenile, pcl 8.7 mm. B, female juvenile, pcl 14.3 mm.

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Pereopod 1 (chelipeds): Unequal; surfaces covered length:height ratio 3.27–4.09 [3.12–3.29] (male), 2.88– with low nodules bearing tufts of golden setae; or- 3.55 [2.59–2.80] (female). Carpus 0.57–0.61 [0.58–0.61] namentation similar on both sides and in both sexes. merus length (male), 0.64–0.71 [0.64–0.67] (female). Coxa and ischiobasis unarmed. Merus outer surfaces Propodus 0.84–0.88 [0.81–0.84] merus length (male), granular; mesial surface relatively smooth, with short 0.90–1.00 [0.89–0.92] (female); length:height ratio ventral spines and strong inner distal spine; dorsal 4.13–5.45 [3.53–3.66] (male), 3.84–4.39 [3.25–3.53] (fe- surface with scattered tubercles, distally with several male). Dactylus 0.86–0.99 [0.86–0.92] propodus length stout spines. Carpus surfaces sparsely tuberculate; (male), 0.88–1.00 [0.84–0.95] (female). dorsal margin with 3 or 4 strong spines. Palm dorsal Pereopod 4 length 2.27–2.49 [2.02–2.21] pcl (male), margin with 3–5 blunt teeth or tubercles, setose; other 2.02–2.26 [1.92–2.03] pcl (female). Merus 0.68–0.76 surfaces smooth apart from setal tufts. [0.65–0.67] pcl (male), 0.56–0.65 [0.55–0.59] pcl (female); Major cheliped 1.73–2.05 [1.68–1.83] pcl (male), length:height ratio 3.23–3.81 [2.97–3.30] (male), 2.99– 1.47–1.63 [1.46–1.51] (female); upper palm length 3.39 [2.72–3.07] (female). Carpus 0.60–0.63 [0.61–0.62] 0.86–1.01 [0.90–1.02] times height (male), 0.87–0.97 merus length (male), 0.66–0.72 [0.67–0.71] (female). [0.92–0.94] (female); occlusal margins of fingers cor- Propodus 0.89–0.94 [0.89–0.91] merus length (male), neous for distal third, proximally with 3 calcareous 0.96–1.20 [0.92–0.97] (female); length:height ratio nodules; dactylus dorsal margin broadly convex, with 4.17–4.91 [3.68–3.97] (male), 3.97–5.50 [3.29–3.70] (fe- rows of tufts of golden setae and blunt proximal tooth, male). Dactylus 0.89–0.95 [0.88–0.93] propodus length 1.23–1.38 [1.22–1.36] times longer than dorsal margin (male), 0.89–1.00 [0.90–0.93] (female). of palm (male), 1.40–1.59 [1.42–1.58] (female). Egg diameter 1.70–2.52 mm. Minor cheliped 1.66–1.82 [1.57–1.72] pcl (male), 1.45–1.61 [1.44–1.49] (female); upper palm length Colour in life. Ivory-white, but often with light brown 0.92–1.28 [0.99–1.04] times height (male), 0.94–1.06 iron precipitates on external cuticle. [1.01–1.07] (female); occlusal margin corneous in distal third to half, proximally crenulate to weakly dentate; Remarks. Paralomis hirtella de Saint Laurent & Macpher- dactylus dorsal margin broadly convex, with rows son, 1997 was first recorded from New Zealand waters of tufts of golden setae and blunt proximal tooth, as P. aff. jamsteci based on specimens from Brothers 1.51–1.86 [1.72–1.85] times longer than dorsal margin Seamount (~35°S) (Webber & Naylor 2004b). Subse- of palm (male), 1.74–1.86 [1.69–1.79] (female). quently, Lörz et al. (2008) and Dawson (2008) reported Pereopods 2–4 (walking legs 1–3): Similar, elongate, P. hirtella from Monowai Caldera (~26°S), just outside covered with low nodules bearing tufts of golden se- of New Zealand waters, including specimens parasit- tae. Pereopod 3 longest. Coxae distal margins smooth, ised by Briarosaccus callosus Boschma. Paralomis hirtella unarmed; ventral surfaces smooth, with scattered setal differs from all other New Zealand lithodids by the tufts. Ischiobasis with low setose nodules, unarmed. smooth, unarmed carapace surface, bearing tufts of Merus extensor margins with 5–8 spines; flexor mar- golden setae and at most a few, low, scattered gran- gins with 3–5 spinose; other surfaces with setal tufts but ules near the anterolateral margins. The characteristic unarmed. Carpus with 2–5 extensor spines. Propodus carapace ornamentation is consistent across the full shorter than merus; with 2–8 short spines on extensor size range examined (pcl 6.7–61.1 mm). margin. Dactylus curved, laterally compressed; longer The specimens from Brothers Seamount agree well than carpus, shorer than propodus; proximally un- with northern specimens (Monowai Caldera material armed; flexor margin lined with 8–12 obliquely inclined and the holotype from the Lau Basin, 22°13’S) in almost corneous spinules; apex corneous. all respects, but differ subtly in having proportion- Pereopod 2 length 2.37–2.61 [2.17–2.29] pcl (male), ally longer chelipeds and walking legs in adults. The 2.05–2.34 pcl [1.97–2.14] (female). Merus 0.75–0.85 smallest ovigerous female from Brothers Seamount is [0.72–0.74] pcl (male), 0.61–0.72 [0.58–0.63] pcl (fe- 35.4 mm pcl, and 29.9 mm pcl from Monowai. In the male); length:height ratio 3.28–3.72 [3.13–3.28] (male), description above, morphometric measurements of 2.94–3.46 [2.63–3.00] (female). Carpus 0.59–0.62 merus the New Zealand specimens are given, followed by length [0.60–0.77] (male), 0.65–0.73 [0.66–0.71] (female). those of the northern specimens in parentheses. Both Propodus 0.82–0.87 [0.81–0.83] merus length (male), Brothers Seamount and northern specimens exhibit 0.88–0.99 [0.86–0.91] (female); length:height ratio typical lithodid sexual dimorphism in which males 3.87–4.74 [3.44–3.76] (male), 3.86–4.20 [3.07–3.41] (fe- have proportionally longer pereopods than respective male). Dactylus 0.85–0.97 [0.74–0.89] propodus length females, differing by about 10% in P. hirtella. Propor- (male), 0.87–0.97 [0.86–0.93] (female). tional pereopod lengths in northern males, however, Pereopod 3 length 2.41–2.63 [2.17–2.31] pcl (male), are similar to those of females from Brothers Seamount. 2.07–2.39 [2.02–2.17] pcl (female). Merus 0.75–0.84 The differences are subtle, and whether they represent [0.71–0.76] pcl (male), 0.62–0.72 [0.60–0.63] pcl (female); phenotypic plasticity or reflect speciation remains to be

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ determined. Note that the pereopod length differences mm, cw 5.8 mm), Pyre Seamount, 42°43.09–43.18’S, between Brothers Seamount and northern specimens 179°54.57–54.87’W, 1075–1008 m, TAN0104/333, 20 were determined independent of the parasitised Apr 2001; NIWA 29642, 1 male (cl 24.0 mm, pcl 19.9 specimens, and thus are not an artefact of parasitic mm, cw 17.8 mm), Zombie Seamount, 42°45.76’S, feminisation. 179°55.51’W, 1019–1081 m, TAN0604/09, 28 May 2006; Six of 23 Monowai Caldera specimens (26%) are NIWA 3916, 1 juvenile female (cl 10.0 mm, pcl 7.8 mm, infected with the rhizocephalan Briarosaccus callosus. cw 6.8 mm), Diabolical Seamount, 42°47.56–47.70’S, Such an infection rate seems disproportionately high 179°58.86–58.60’W, 950–900 m, TAN0104/47, 16 Apr in comparison to other New Zealand lithodids. For 2001. instance, of the 83 specimens of Lithodes aotearoa examined, only four specimens (5%) were obviously infected Other material examined. Western Australia: NMV (i.e., externae visible). The infected males of P. hirtella J54984, 1 juvenile female (cl 19.0 mm, pcl 14.4.mm, have been feminised, their pereopod proportions being cw 12.5 mm), off Albany, 35°26.05’S, 118°21.00’E, similar to those of sympatric females. Saint Laurent & 900–915 m, RV Southern Surveyor, SS1005/44/040, 25 Macpherson (1997) also recorded two parasitised males Nov 2005. among the 10 specimens examined. It is noteworthy Tasmania: NMV J44012, ovigerous female (cl 32.4 that no southern specimens exhibited any evidence of mm, pcl 27.3 mm, cw 23.3 mm), 83.8 km SSE of South- parasitism, either in visible rhizocephalan externae or east Cape, “J1” seamount, 44°16.2’S, 147°19.6’E, 987 morphometric anomalies, despite the vents at Brothers m, epibenthic sled, RV Southern Surveyor, SS01/97/36, Seamount being within the known range latitudinal coll. T.N. Stranks et al., 27 Jan 1997. and bathymetric range of Briarosaccus callosus. Paralomis hirtella is a hydrothermal vent associate, Diagnosis. Carapace pyriform, slightly longer than and appears to be common around vents on Brothers wide, margins and surface covered with upright spines, Seamount, where other vent decapods have also been surfaces minutely setose; outer distance between bases recorded: Alvinocaris alexander Ahyong, 2009, A. niwa of anterolateral spines exceeding half carapace width. Webber, 2004, A. longirostris Kikuchi & Ohta, 1995, Rostrum with 2 pairs of dorsal spines; subrostral lobe Nautilocaris saintlaurentae Komai & Segonzac, 2004, rounded. Dorsal eyespines not extending anteriorly Lebbeus wera Ahyong, 2009, and Gandalfus puia McLay, beyond cornea. Scaphocerite multispinose. Abdominal 2007 (McLay 2007; Ahyong 2009). somite 2 and 3 spinose; somites 4–6 with tubercles. Chelipeds and walking legs covered with spines, most Distribution. North Fiji Back-Arc Basin (White Lady elongate on dorsal and extensor margins; spines on vent field); Lau Basin Back-Arc Basin (Hina Hina vent dactylar extensor margin restricted to proximal half field), Monowai Caldera, and from Brothers Seamount, of margin. New Zealand; 1041–1750 m. Description. Carapace: Pyriform, 1.07–1.17 times longer Paralomis poorei sp. nov. than wide; regions distinct; surface and margins uniformly covered with slender, straight conical spines (Figs 81, 92–97, Pl. 3E) of varying length, 7 gastric, 4 cardiac and 4 branchial Paralomis cf. phrixa. — Koslow & Gowlett-Holmes, 1998: 43. longer than other spines; surface of spines with short, — Poore, 2004: 269, fig. 76c. — Poore et al., 2008: 27. well-spaced simple setae; longest spine (on anterior Paralomis sp. MoV2717. — Poore et al., 1998: 66, 72. branchial margin) about 0.2 pcl; cardiac region with 20–23 spines; cervical groove distinct. Pterygosto- Type material. (All Chatham Rise, New Zealand). mian region sparsely granulose; with prominent, Holotype: NIWA 61161, male (cl 24.6 mm, pcl 18.9 mm, anterior, submarginal spine; spinulose on posterior cw 17.3 mm), Diabolical Seamount, 42°47.17–46.96’S, two-thirds. 179°59.11–59.01’W, 993–900 m, TAN0104/48, 16 Apr Rostrum 0.19–0.32 pcl; broad basally, not constricted 2001. proximal to dorsal spines; median spine slender, ven- Paratypes: NIWA 53273, 1 juvenile male (cl 11.8 mm, tral lobe angular, unarmed; proximally with 2 pairs of pcl 9.1 mm, cw 7.2 mm), 42°38.65–38.67’S, 179°52.88– laterally divergent spines directed obliquely upwards 53.05’E, 1052–1080 m, TAN0905/48, 18 Jun 2009; NIWA and single upright spine between proximal pair of dor- 29526, 1 juvenile male (cl 14.9 mm, pcl 11.3 mm, cw sal spines. Posterior orbital margin concave, unarmed; 10.1 mm), Pyre Seamount, 42°43.06’S, 179°54.29’W, outer orbital spine slender, directed anteriorly, not 1025–1156 m, TAN0604/102, 4 Jun 2006; NIWA 53134, reaching beyond apex of cornea (when eyes directed 1 male (pcl 27.3 mm, cw 25.4 mm), 42°44.68–44.90’S, anteriorly). Anterolateral spine as long as outer orbital 179°55.41–55.43’E, 1020–1125 m, TAN0905/41, 17 Jun spine; outer distance between bases of anterolateral 2009; NIWA 3917, 1 juvenile male (cl 9.0 mm, pcl 6.7 spines exceeding half carapace width.

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 92. Paralomis poorei sp. nov., male holotype, cl 24.6 mm, pcl 18.9 mm, cw 17.3 mm, Diabolical Seamount (NIWA 61161). A, dorsal habitus. B, carapace, right lateral view.

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 93. Paralomis poorei sp. nov., male holotype, cl 24.6 mm, pcl 18.9 mm, cw 17.3 mm, Diabolical Seamount (NIWA 61161). A, right chela, outer surface. B–C, left chela, outer and inner surfaces. D–E, right pereopod 4, dorsal and ventral view. F, anterior carapace, right oblique view.

Ocular peduncle: Longer than cornea; dorsally with length of distal antennular peduncle article. 5 or 6 acute granules and single short distal spine; Antenna: Basal antennal article with short ante- distal spine upright, about half corneal diameter but rolateral spine. Article 2 unarmed on inner margin; not overreaching cornea. outer margin with acute granule proximally and long Antennule: Peduncle unarmed, reaching anteriorly distal spine reaching slightly beyond apex of article 3. beyond apex of antennal peduncle by half to quarter Article 3 unarmed. Scaphocerite a long, slender spine

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 94. Paralomis poorei sp. nov., male holotype, cl 24.6 mm, pcl 18.9 mm, cw 17.3 mm, Diabolical Seamount (NIWA 61161). A, carapace. B, anterior carapace, dorsal view. C, abdomen. D, posterior carapace and abdominal somite 2.

overreaching distal peduncular article; with 2 lateral tubercles and some spines of about half length of spines spines, first minute, second about two-thirds length of on preceding somite. Somites 4–5 sparsely granulate, main spine; dorsally unarmed; mesially with 2 small margins irregularly granulate. Somite 6 with 4 blunt, spines or acute granules proximally. Article 4 unarmed, apically setose tubercles on distal margin. Telson semi- almost half length of article 5. circular, unarmed. Abdomen: Ornamentation of both sexes similar. Pereopod 1 (chelipeds): Strongly spinose, unequal. Somite 2 covered with spines of similar length to dorsal Coxae unarmed, setose; distal margins with dense tufts carapace spines. Somite 3 less spinose, with scattered of setae. Ischiobasis spinose laterally, ventrally with

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 95. Paralomis poorei sp. nov., male holotype, cl 24.6 mm, pcl 18.9 mm, cw 17.3 mm, Diabolical Seamount (NIWA 61161). A, right antenna, lateral view. B, right antenna, dorsal view. C, rostrum, right lateral view. D, right eye, lateral view. Scale = 1.0 mm.

clusters of golden setae. Merus with smooth mesial tate. Ischiobasis spinose. Merus ovate in cross section; surface, others surfaces spinose, long prominent spine extensor and flexor margins each with 2 relatively dorsodistally. Carpus prominently spinose except for even rows of 7–11 and 5–7 spines respectively; dorsal mesial surface, with clusters of golden setae. Palm surface with smaller scattered spines; ventral surface outer surface with 2 or 3 uneven rows of 4–6 spines; unarmed. Carpus spinose dorsally and laterally, long- upper margin with 2 uneven rows of 4–7 spines; ven- est spine as high as carpus. Propodus dorsoventrally tral surface irregularly spinose or tuberculate; mesial flattened; extensor margin with 2 rows of 6–8 spines; surface with blunt scattered tubercles bearing tufts of flexor margin with row of 7–9 spines; dorsal surface golden setae. multispinose; ventral spines sparse and short, not ex- Major cheliped 1.61–2.10 pcl (male), 1.49 pcl (fe- ceeding 1.5 length of dorsal spines. Dactylus curved, male); upper palm length 1.19–1.29 times height (male), laterally compressed; longer than carpus; with 5–7 1.31 (female); occlusal margins of fingers corneous for spines on proximal half and scattered tufts of golden distal quarter to fifth, proximally with 3 calcareous setae; flexor margin lined with 13–18 obliquely inclined nodules; dactylus dorsal margin broadly convex, with corneous spinules; apex corneous. rows of tufts of golden setae and small proximal spine, Pereopod 2 length 1.90–2.31 pcl (male), 1.69–1.81 1.06–1.12 times longer than dorsal margin of palm pcl (female). Merus 0.62–0.74 pcl (male), 0.51–0.56 pcl (male), 1.09 (female). (female); length:height ratio 4.22–4.61 (male), 3.60–3.97 Minor cheliped 1.54–1.93 pcl (male), 1.49 pcl (fe- (female). Carpus 0.60–0.61 merus length (male), 0.64– male); upper palm length 1.33–1.48 times height (male), 0.70 (female). Propodus 0.85–0.87 merus length (male), 1.39 (female); occlusal margin corneous in distal third 0.94–0.98 (female); length:height ratio 5.38–6.82 (male), to half, proximally crenulate; dactylus dorsal margin 5.11–6.30 (female). Dactylus 0.76–0.77 propodus length broadly convex, with rows of tufts of golden setae (male), 0.75–0.79 (female). and small proximal spine, 1.62–1.65 times longer than Pereopod 3 length 1.91–2.26 pcl (male), 1.72–1.79 pcl dorsal margin of palm (male), 1.59 (female). (female). Merus 0.59–0.69 pcl (male), 0.54 pcl (female); Pereopods 2–4 (walking legs 1–3): Similar, elongate, length:height ratio 4.00–4.65 (male), 3.59–3.90 (female). spinose. Pereopod 2 longest. Coxae unarmed, or with Carpus 0.61 merus length (male), 0.66–0.68 (female). a few low granules, distal margins crenulate to den- Propodus 0.93–0.94 merus length (male), 0.90–0.99

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 96. Paralomis poorei sp. nov., ovigerous female, cl 32.4 mm, pcl 27.3 mm, cw 23.3 mm, J1 seamount (NMV J44012). A, dorsal habitus. B, carapace. C, abdomen. E, right chela.

(female); length:height ratio 5.88–7.15 (male), 5.01–6.55 5.38–6.60 (female). Dactylus 0.73–0.80 propodus length (female). Dactylus 0.75–0.81 propodus length (male), (male), 0.73–0.79 (female). 0.79–0.82 (female). Pereopod 4 length 1.81–2.17 pcl (male), 1.60–1.80 Colour in life. Translucent pink-orange overall (Pl. pcl (female). Merus 0.53–0.64 pcl (male), 0.46–0.49 pcl 3E). (female); length:height ratio 3.71–4.37 (male), 3.32–3.88 (female). Carpus 0.66–0.68 merus length (male), 0.72– Etymology. Named after Gary Poore, who first re- 0.74 (female). Propodus 1.00–1.04 merus length (male), ported the presence of the species from southeastern 1.06–1.07 (female); length:height ratio 5.77–7.00 (male), Australia.

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 97. Paralomis poorei sp. nov. A–B, male paratype, pcl 27.3 mm, cw 25.4 mm (NIWA 53134), dorsal habitus and chelipeds. C, juvenile male, pcl 11.3 mm (NIWA 29526). D, juvenile male paratype, pcl 6.7 mm (NIWA 3917).

Remarks. Paralomis poorei sp. nov. most closely resem- maturity not known for P. makarovi). Paralomis poorei bles P. bouvieri Hansen, 1908 [North Atlantic] and P. differs from P. bouvieri in the presence of a subrostral makarovi Hall & Thatje, 2009b [Bering Sea, North Pa- lobe; the distal eyespine does not overreach the cornea; cific] in carapace physiognomy and spination, and in and the ventral surfaces of the meri of the walking the similar armature and length of the walking legs. legs are smooth or sparsely granulate rather than Each is apparently a small species, with P. poorei and spinose. Paralomis poorei is readily distinguished from P. bouvieri maturing at less than 30 mm pcl (size at P. makarovi in the presence of the subrostral lobe and in

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ the length of the setae on the surfaces of the carapace almost as long as the initial ‘long’ spines. These long spines — markedly shorter, rather than markedly spines are homologous with the short dorsal spines longer than the basal width of the spines. of comparatively less ornamented species such as Of the New Zealand and Australian Paralomis spe- P. birsteini Macpherson, 1988a, P. gowlettholmes sp. nov., cies, P. poorei could be confused with P. webberi sp. and P. stevensi Ahyong & Dawson, 2006. nov. (New Zealand), juvenile P. zealandica Dawson The Australian and New Zealand specimens of & Yaldwyn, 1971 (New Zealand), P. echidna sp. nov. P. poorei agree well. Differences, namely in proportional (Australia), and P. taylorae sp. nov. (Australia) owing limb lengths, are consistent with sexual dimorphism to their dense dorsal spination. The tuberculate rather — the New Zealand specimens are all males and both than spinulate abdominal somites will distinguish Australian specimens are female. Therefore, detailed P. poorei from each of the aforementioned species. comparison of both sexes from both regions is not yet Paralomis poorei further differs from P. webberi, possible, but differences between New Zealand and P. echidna, and size-matched P. zealandica in having Australian populations might become evident when more elongate, more slender walking legs as meas- more material becomes available for study. ured by pereopod 4 length and pereopod 4 meral Paralomis poorei is the smallest known lithodid in proportions, and from P. taylorae in having shorter, less New Zealand and Australian waters (females oviger- slender walking legs (see Tables 4, 5). The setation of ous at less than 30 mm pcl) and is one of only four the carapace spines also differs among species, being lithodid species to be shared by both regions (see also at most apically setose in P. zealandica and glabrous in P. echidna sp. nov., P. staplesi sp. nov., and Lithodes P. taylorae, rather than sparsely scattered over the spine macquariae sp. nov.). Species previously thought to surface as in P. webberi, P. echidna, and P. poorei. occur on both sides of the Tasman are, as a result of The juveniles agree with adults in having similar the present study, now known to be composites (see abdominal and pereopodal ornamentation, and dis- Lithodes aotearoa sp. nov., L. australiensis sp. nov., Neo- tinctly unequal chelipeds (in males). The carapace lithodes brodiei Dawson & Yaldwyn, 1970, N. flindersi ornamention is largely consistent with adults except Ahyong, 2010). in the smallest juveniles (pcl 6.7–9.1 mm) in which the spination of the pereopods and carapace surface is less Distribution. Presently known from seamounts on extensive. The carapace surface of the small juveniles the Chatham Rise, New Zealand, from seamounts off is armed with long dorsal spines (seven gastric, four southeastern Tasmania, and southwestern Australia; cardiac, and four branchial) and numerous minute 900–1156 m. spines or acute granules which, in adults, become

Table 4. Selected measurements of male P. poorei, P. echidna, P. webberi, P. taylorae, and juvenile P. zealandica. Size range of P. zealandica selected to correspond to known size range of P. poorei. Abbreviations: postorbital carapace length (pcl); pereopod 4 (P4); length (L); height (H).

Size range P4 L/pcl P4 merus L/pcl P4 merus L/H P4 propodus (pcl) (Male) (Male) (Male) L/H (Male) P. poorei 11.3–27.3 1.81–2.17 0.53–0.64 3.71–4.37 5.40–7.00 P. echidna 49.7 1.56 0.48 2.91 4.34 P. webberi ? ? ? ? ? P. zealandica 9.0–25.7 1.44–1.53 0.43–0.48 2.74–3.01 3.61–4.31 P. taylorae ? ? ? ? ?

Table 5. Selected measurements of female P. poorei, P. echidna, P. webberi, P. taylorae, and juvenile P. zealandica. Size range of P. zealandica selected to correspond to known size range of P. poorei. Abbreviations: postorbital carapace length (pcl); pereopod 4 (P4); length (L); height (H).

Size range P4 L/pcl P4 merus L/pcl P4 merus L/H P4 propodus (pcl) (Female) (Female) (Female) L/H (Female) P. poorei 14.4–27.3 1.60–1.80 0.46–0.51 3.32–3.88 5.38–6.60 P. echidna 33.6–38.6 1.20–1.28 0.35–0.37 2.43–2.52 3.68–3.73 P. webberi 33.6 1.23 0.36 2.47 4.67 P. zealandica 8.9 1.50 0.44 2.68 3.82 P. taylorae 39.8 2.74 0.85 4.35 10.6

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Paralomis staplesi sp. nov. Antennule: Peduncle unarmed, reaching anteriorly (Figs 81, 98–100, Pl. 3H) beyond antennal peduncle by half-length of distal antennular peduncle article. Type material. Holotype: NMV J61052, male (cl 59.1 mm, Antenna: Basal antennal article with small anterola- pcl 50.0 mm, cw 49.0 mm), Tasman Fracture Zone, teral spine. Article 2 with inner distal spine; outer mar- Tasmania, 45°18’01’S, 146°07’14”E, 2213 m, FV Tommy gin with minute basal spine and slender distal spine Thompson, TN228-J2-392-012-005, ROV Jason 2, coll. D. reaching slightly beyond article 4. Article 3 unarmed. Staples, 11 Jun 2009. Scaphocerite slender, reaching beyond midlength of article 5, with 1 or 2 short inner spines and 2 outer spines. Other material examined. NIWA 3920, 1 badly frag- Article 4 unarmed, about half length of article 5. mented male, Kermadec Ridge, 32°32.334–33.551’S, Abdomen of male: Somite 2 with small, widely sepa- 179°39.616–38.977’W, 2312–1958 m, TAN0205/82, 25 rated spines along margins and on surface. Surface of Apr 2002. somites 3–5 sparsely granular or tuberculate. Somite 3 median plate unarmed; submedian plates with low, Diagnosis. Carapace subpentagonal, length and width irregular, triangular teeth on lateral margin; marginal subequal; surface glabrous, covered with small, scat- plates absent, apparently undifferentiated. Somites tered granules and numerous, small, well-spaced coni- 4–5 with unarmed median and submedian plates; cal spines in addition to larger, longer spines (including marginal plates undivided, with irregular triangular 5 gastric, 4 cardiac, 4 branchial); lateral margins lined teeth. Somite 6 slightly wider than long, subpentagonal, with well-spaced spines; outer distance between bases with pair of small distal teeth; marginal plates short, of anterolateral spines half carapace width. Rostrum triangular. Telson wider than long, rounded. trispinose, broadest basally. Scaphocerite with inner Pereopod 1 (chelipeds): Spination similar in both sexes. and outer spines. Chelipeds and walking legs strongly Major cheliped of males inflated, 1.59 times height of spinose. Walking legs elongate, about 3 times pcl; dac- minor cheliped; minor cheliped slender. Coxa smooth, tyli shorter than extensor margin of propodi. Walking unarmed; distal margins with dense tufts of setae. Is- leg 3 merus about as long as pcl, length exceeding 6.7 chiobasis with 8 stout, apically setose ventral spines. times height (males); propodus length exceeding 9 Merus with smooth mesial margins and tuberculate times height. lateral margins; dorsal and ventral margins spinose, inner distal spine largest. Carpus with prominent Description of holotype. Carapace: Subpentagonal, 1.02 spines on dorsal and lateral surfaces; dorsal margin times longer than wider; surface covered with small, with rows of 3–5 spines; mesial margin with 2 spines, scattered granules and numerous, well-spaced conical proximal largest; ventral surface with 3 short spines. spines, glabrous. Gastric region convex, elevated, more Palm mesial margin with 6 spines, other surfaces with prominent than other regions; in addition to scattered prominent, apically setose tubercles or acute tubercles. spines, with 5 longer conical spines forming pentagon Fingers with short basal tubercle and rows of tufts of with anteriormost in midline. Lateral margin of he- golden setae. patic region with 3 conical spines, anterior 2 shorter, Major cheliped 1.74 pcl (male); upper palm length posterior long, curved. Branchial regions with 13 or 1.17 times height (male); occlusal margins of fingers 14 prominent conical marginal spines; surface with corneous for distal third, proximally with 3 calcareous 4 spines distinctly longer than other surface spines. nodules; dactylus dorsal margin broadly convex, with Cardiac region subtriangular, with 2 pairs of short coni- rows of tufts of golden setae and blunt proximal tooth, cal spines forming square. Intestinal region sparsely 1.21 times longer than dorsal margin of palm (male). spinose. Pterygostomian region sparsely granular, with Minor cheliped 1.41 pcl (male); upper palm length prominent anterior spine. 1.27 times height (male); occlusal margin corneous for Rostrum 0.18 pcl; broadest basally, without con- slightly less than distal third, proximally crenulate; striction; median spine smooth ventrally; paired dactylus dorsal margin broadly convex, with rows of dorsal spines divergent, directed obliquely upwards. tufts of golden setae and blunt proximal tooth, 1.81 Posterior orbital margin slightly concave; outer orbital times longer than dorsal margin of palm (male). spine reaching to end of cornea. Anterolateral spine as Pereopods 2–4 (walking legs 1–3): Similar, elongate, long as outer orbital spine, with short marginal spinule spinose. Coxae sparsely granular; distal margins with midway between outer orbital spine and anterolateral 3 blunt apically setose teeth. Ischiobasis with 3–5 api- spine; outer distance between bases of anterolateral cally setose ventral spines. Merus slightly compressed, spines half carapace width. shorter than carapace (ischiomerus longer than cara- Ocular peduncle: Longer than cornea; with 3 acute pace); extensor margin with 11–13 spines in addition to dorsal granules and distal spine slightly overhanging paired distal spines; dorsal surface with 10–14 spines; cornea. flexor margin with 2 uneven rows of 5–7 spines; ventral

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 98. Paralomis staplesi sp. nov., male holotype, cl 59.1 mm, pcl 50.0 mm, cw 49.0 mm, Tasman Fracture Zone, Tasmania (NMV J61052). A, dorsal habitus. B, carapace, right lateral view.

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 99. Paralomis staplesi sp. nov., male holotype, cl 59.1 mm, pcl 50.0 mm, cw 49.0 mm, Tasman Fracture Zone, Tasmania (NMV J61052). A, carapace, B, right pereopod 4.

surface with scattered granules; merus of pereopod 3 Pereopod 4 length 2.96 pcl (male). Merus 0.95 pcl slightly longer than that of pereopod 2 and pereopod 4. (male); length:height ratio 6.75 (male). Carpus 0.51 Carpus half merus length, subcircular in cross section; merus length (male). Propodus 0.89 merus length extensor margin with 9 spines of which 3 or 4 mark- (male); length:height ratio 9.81 (male). Dactylus 0.80 edly longest; dorsal surface with 6–8 spines; ventral propodus length (male). surface with 3–5 small spines; flexor margin unarmed. Propodus dorsoventrally flattened; slightly shorter Colour in life. Overall purplish-red (Pl. 3H). than merus; dorsal surface, extensor and flexor margins prominently multispinose. Dactylus broadly curved; Etymology. Named for David Staples, Museum Victo- slightly shorter than extensor margin of propodus; ria, who collected the holotype. surface with tufts of setae; extensor margin with 3 apically corneous spines proximally; lateral proximal Remarks. Paralomis staplesi sp. nov. closely resembles surfaces with short, distinct sulcus, flanked ventrally P. birsteini Macpherson, 1988a, P. stevensi Ahyong & by small, corneous spine; flexor margin lined with Dawson, 2006, and P. gowlettholmes sp. nov. in overall corneous spinules carapace shape, structure, and ornamentation of the Pereopod 2 length 2.89 pcl (male). Merus 0.98 pcl pereopods. The new species differs primarily in hav- (male); length:height ratio 6.90 (male). Carpus 0.49 ing more prominent and more numerous carapace and merus length (male). Propodus 0.81 merus length pereopodal spination (compare Figs 63, 83, 102, 106). In (male); length:height ratio 9.02 (male). Dactylus 0.84 P. birsteini, P. stevensi, and P. gowlettholmes, the carapace propodus length (male). is sparsely tuberculate or granulate, with several small Pereopod 3 length 3.03 pcl (male). Merus 1.02 pcl conical spines on the gastric, hepatic, and branchial (male); length:height ratio 7.39 (male). Carpus 0.48 regions. The dorsal carapace tubercles or granules merus length (male). Propodus 0.83 merus length and small spines of the aforementioned species are (male); length:height ratio 9.84 (male). Dactylus 0.82 developed in P. staplesi as small but prominent spines propodus length (male). and large, prominent, conical spines, respectively.

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 100. Paralomis staplesi sp. nov., male holotype, cl 59.1 mm, pcl 50.0 mm, cw 49.0 mm, Tasman Fracture Zone, Tasmania (NMV J61052). A, right chela. B, left chela. C, posterior carapace and abdominal somite 2. D, ventral surface and abdomen. E, anterior carapace, dorsal view. F, right orbit and antenna.

Similarly, the spines of the walking legs of P. staplesi more elongate walking legs in which the pereopod 4 are more numerous and proportionally longer than in meri are as long as or longer than the carapace (shorter P. birsteini, P. stevensi, and P. gowlettholmes. In addition, in P. birsteini and P. gowlettholmes), and in which the Paralomis staplesi resembles P. stevensi and differs from walking leg dactyli are shorter than the propodal ex- P. gowlettholmes and P. birsteini in its more slender, tensor margins.

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ The male specimen from the Kermadec Ridge is (cl 68.4 mm, pcl 57.2 mm, cw 58.2 mm), area 88.2E, badly fragmented: the abdomen, chelipeds and most 69°41.00–40.00’S, 124°31.00–38.00’W, 1432–1560 m, FV walking legs are largely intact but the carapace is San Liberatore, trip 1743, set 78, sample 12, camera 1, crushed with only portions of the right branchial mar- photo 18, coll. G. Dolan, 11 Apr 2003. gin, the left anterolateral margin and the rostral/orbital regions remaining. In all details that can be compared, Diagnosis. Carapace subpentagonal, length and width the damaged specimen agrees well with the holotype. subequal; surface glabrous, sparsely covered with A complete specimen may reveal specific differences, small, scattered granules and short conical spines but until further material becomes available, it is including median gastric spine; lateral margins with tentatively assigned to P. staplesi. Paramonis staplesi, short conical spines; outer distance between bases of P. echidna sp. nov., and P. poorei sp. nov. are the only anterolateral spines half carapace width. Rostrum species of Paralomis known from both New Zealand trispinose, broadest basally, without constriction. and Australian waters. Scaphocerite with inner and outer spines. Mature male chelipeds strongly dimorphic, spinose; major Distribution. Presently known only from Tasmania and cheliped palm 1.7–1.9 times height of minor cheliped. the Kermadec Ridge, New Zealand; 1958– 2312 m. Major cheliped palm of female about 1.3 times height of minor cheliped. Walking legs elongate, 2–3 times Paralomis stevensi Ahyong & Dawson, 2006 pcl, spinose; ventral spines of propodi widely spaced; dactyli shorter than extensor margin of propodi. Walk- (Figs 66, 101–104, Pl. 3G) ing leg 3 merus as long as or longer than pcl (male), Paralomis birsteini. — Stevens, 2004: 6, 7 [not P. birsteini about 0.6 pcl (female), length exceeding 6.5 times height Macpherson, 1988]. (males), 4.7 (female); propodus length exceeding 8.4 Neolithodes brodiei. — Thatje & Lörz, 2005: 335 [NIWA 3435 times height (male), 6.7 (females). only; not N. brodiei Dawson & Yaldwyn, 1970]. Paralomis stevensi Ahyong & Dawson, 2006: 59–66, figs 7–10 Description. Carapace: Subpentagonal, length 0.97–1.04 [type locality: near Scott Island, Ross Sea, 65°40.0–36.0’S, times width; surface glabrous, sparsely covered with 176°21.0–17.0’W, 1688–1924 m]. small, scattered granules. Gastric region convex, elevated, more prominent than other regions, with 5 short Type material. Holotype: NIWA 23843, male (cl 84.0 mm, conical spines forming pentagon with anteriormost in pcl 74.6 mm, cw 75.7 mm), near Scott Island, 65°40.0– midline. Lateral margin of hepatic region with 3 short 36.0’S, 176°21.0–17.0’W, 1688–1924 m, from stomach of conical spines, anterior shorter. Branchial regions with Dissostichus mawsoni Norman, RV Avro Chieftain, set 3, 12–14 short marginal spines; surface with 4 or 5 low coll. D. Stevens, 2 Dec 2003. conical spines in addition to low, scattered tubercles. Paratypes: NIWA 3435, 1 ovigerous female (badly Cardiac region subtriangular, with 2 pairs of short coni- damaged, pcl about 61.2 mm), E of Sturge Island, cal spines forming square. Intestinal region sparsely Balleny Islands, 67°25.37’S, 165°15.82’E, 1389 m, granular. Pterygostomian region sparsely granular, TAN0402/214, 3 Mar 2004; MNZ Cr10856, 1 male (cl with prominent anterior spine. 73.4 mm, pcl 61.7 mm, cw 62.0 mm), Ross Sea, 66°46.90– Rostrum 0.13–0.25 pcl; broadest basally, without 44.90’S, 172°56.4–173°01.70’W, 1846–1420 m, found on constriction; median spine smooth ventrally; paired hook, RV Avro Chieftain, trip 1742, set 51, coll. J. Buirski, dorsal spines divergent or subparallel, directed ob- 14 Mar 2003. liquely upwards. Posterior orbital margin near concave; outer orbital spine reaching to base of cornea. Other material examined. Ross Sea, Antarctica: NMNZ Anterolateral spine as long as or shorter than outer Cr11743–11744, 1 male (cl 82.4 mm, pcl 71.0 mm, cw orbital spine, with low marginal granule or small spine 70.2 mm), 1 ovigerous female (cl 87.8 mm, pcl 70.4 midway between outer orbital spine and anterolateral mm, cw 67.9 mm), 64°41.00–44.7’S, 172°59.7–55.2’W, spine; outer distance between bases of anterolateral 1327–1163 m, Trip 1742, set 84, FV Avro Chieftain, spines half carapace width. coll. G. Anderson, 7 Apr 2003; NMNZ Cr11254, 1 Ocular peduncle: Longer than cornea; with 2–5 dorsal parasitised male (cl 88.5 mm, pcl 75.0 mm, cw 76.0 granules. mm), 66°35’S, 177°33’W, 1549 m, trip 1728, set 40, FV Antennule: Peduncle unarmed, reaching anteriorly Janas; NIWA 38499, 1 walking leg, 67°24.48–24.33’S, beyond antennal peduncle by about three-quarters 179°48.55–48.28’W, 1520–1560 m, TAN0802/237, 7 Mar length of distal antennular peduncle article. 2008; NIWA 27835, 1 male (cl 96.7 mm, pcl 81.7 mm, cw Antenna: Basal antennal article with small ante- 84.6 mm), 70°49’S, 178°41’W, trip 2333, set 47, FV San rolateral spine. Article 2 with angular to sharp inner Aspiring, coll. Jano van Heerden, 17 Jan 2007. distal margin; outer margin with small basal spine and Amundsen Sea, Antarctica: NMNZ Cr11804, 1 male slender spine not reaching beyond article 4. Article 3

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 101. Paralomis stevensi Ahyong & Dawson, 2006, male, pcl 71.0 mm, Ross Sea, Antarctica (MNMZ Cr11743). A, dorsal habitus. B, carapace. C, ventral surface and abdomen.

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 102. Paralomis stevensi Ahyong & Dawson, 2006, male, pcl 71.0 mm, Ross Sea, Antarctica (MNMZ Cr11743). A, right chela. B, posterior carapace and abdominal somite 2. C, left chela. D, right pereopod 4. E, carapace, right lateral view. F, anterior carapace, dorsal view.

unarmed. Scaphocerite slender, reaching beyond mi- unarmed median and submedian plates; marginal dlength but not beyond distal three-quarters of article plates undivided, with 2–4 low angular protrusions. 5, with 1 or 2 short inner spines and 1 or 2 outer spines. Somite 6 as long as or longer than wide, subquadrate, Article 4 unarmed, about half length of article 5. with pair of small distal teeth; marginal plates short, Abdomen: Ornamentation similar in both sexes. triangular. Telson wider than long, apex rounded. Somites sparsely granular. Somite 2 with small, widely Pereopod 1 (chelipeds): Spination similar in both sexes. separated spines along margins. Somite 3 median Major cheliped of males strongly inflated, 1.66–1.91 plate unarmed; submedian plates with low, irregular, times height of minor cheliped; minor cheliped slender. triangular teeth on lateral margin; marginal plates Chelipeds of female unequal, major cheliped 1.31–1.33 absent, apparently undifferentiated. Somites 4–5 with times height of minor cheliped. Coxae smooth, un-

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 103. Paralomis stevensi Ahyong & Dawson, 2006, male, pcl 81.7 mm, Ross Sea, (NIWA 27835). A, carapace. B, chelipeds. C, anterior carapace, dorsal view. D, right pereopod 4 dactylus and propodus.

armed; distal margins with dense tufts of setae. Is- Major cheliped 1.62–2.22 pcl (male), 1.32–1.36 chiobasis with 4–8 stout, apically setose ventral spines. (female); upper palm length 1.03–1.15 times height Merus with smooth mesial margins and tuberculate lat- (male), 1.19–1.24 (female); occlusal margins of fingers eral margins; dorsal and ventral margins spinose, inner corneous for distal third, proximally with 3 calcareous distal spine largest. Carpus with prominent spines on nodules; dactylus dorsal margin broadly convex, with dorsal and lateral surfaces; dorsal margin with row of rows of tufts of golden setae and blunt proximal tooth, 3–5 spines; mesial margin with 2 spines, proximal larg- 1.04–1.15 times longer than dorsal margin of palm est; ventral surface with 3 spines. Palm mesial margin (male), 1.26–1.28 (female). with 6 spines, other surfaces with prominent, apically Minor cheliped 1.53–1.86 pcl (male), 1.31–1.39 setose tubercles or acute tubercles. Fingers with short (female); upper palm length 1.15–1.31 times height basal tubercle and rows of tufts of golden setae. (male), 1.28–1.32 (female); occlusal margin corneous

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 104. Paralomis stevensi Ahyong & Dawson, 2006, ovigerous female, pcl 70.4 mm, Ross Sea (NMNZ Cr11744). A, dorsal habitus. B, carapace. C, ventral surface and abdomen.

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ for slightly less than distal third, proximally crenulate; the Ross Sea, Antarctica. The availability of additional dactylus dorsal margin broadly convex, with rows of specimens, including an intact ovigerous female, has tufts of golden setae and blunt proximal tooth, 1.57– permitted more complete documentation of the species; 1.68 times longer than dorsal margin of palm (male), a redescription is provided above. 1.55–1.88 (female). Paralomis stevensi most closely resembles P. birst- Pereopods 2–4 (walking legs 1–3): Similar, elongate, eini Macpherson, 1988a (type locality: Ross Sea) and spinose. Coxae smooth, unarmed; distal margins with P. gowlettholmes sp. nov. (type locality: Tasmania) dense tufts of setae. Ischiobasis with 3–5 apically se- in the combination of an armed inner margin of the tose ventral spines. Merus compressed, shorter than scaphocerite and carapace ornamentation in which carapace in both sexes; extensor margin with 9–11 the primary spines are low and simple, and the sur- spines in addition to paired distal spines; dorsal surface face between the primary spines is smooth or sparsely with 5–8 spines; flexor margin with 2 rows of 4 or 5 granular. In addition to reaching a larger overall size, spines; merus of pereopod 3 slightly longer than that P. stevensi can be immediately distinguished from of pereopod 2 and pereopod 4. Carpus slightly longer P. gowlettholmes by the proportionally shorter walk- than half merus length, subcircular in cross section; ing leg dactyli — shorter, rather than longer than the extensor margin with 7 or 8 spines; dorsal surface with extensor margin of the respective propodi. Paralomis 7 or 8 spines; flexor margin unarmed. Propodus dors- stevensi chiefly differs fromP . birsteini in the following oventrally flattened; slightly shorter than merus; with features: the dactyli of the walking legs are shorter, 11–13 spines on extensor margin; dorsal surface with rather than slightly longer than the extensor margin small scattered spines; flexor margin with 7 or 8 spines. of the propodi; the right cheliped of large males (pcl Dactylus broadly curved; slightly shorter than exten- 71.0–81.7 mm) is considerably more strongly inflated sor margin of propodus; surface with tufts of setae; than the left, with the palm 1.8–1.9 times the height of extensor margin with 3 or 4 apically corneous spines the left, in contrast to 1.5–1.6 for P. birsteini in a similar proximally; lateral proximal surfaces with short, dis- size range (pcl 69.5–90.9 mm). The right palm of female tinct sulcus, flanked ventrally by 1 or 2 small, corneous P. stevensi exceeds 1.3 times the height of the left com- spines; flexor margin lined with corneous spinules. pared to less than 1.3 (1.24–1.25) in P. birsteini. As also Pereopod 2 length 2.63–3.24 pcl (male), 2.06–2.10 indicated by Ahyong & Dawson (2006), the walking pcl (female). Merus 0.86–1.10 pcl (male), 0.62–0.68 pcl legs of male P. stevensi are proportionally longer and (female); length:height ratio 5.82–7.35 (male), 4.17–4.52 more slender than in P. birsteini of similar size. Com- (female). Carpus 0.50–0.55 merus length (male), 0.60 paring large, similarly sized males of P. stevensi (pcl (female). Propodus 0.83–0.86 merus length (male), 81.7 mm) and P. birsteini (pcl 79.4 mm), the pereopod 0.89–0.94 (female); length:height ratio 7.50–9.22 (male), 4 length is 3.3 versus 2.7 pcl, the pereopod 4 merus 6.12–7.39 (female). Dactylus 0.76–0.89 propodus length length:height ratio is 7.0 versus 4.9, and the pereopod (male), 0.80–0.86 (female). 4 propodus length:height ratio is 10.7 versus 6.7. Note Pereopod 3 length 2.80–3.33 pcl (male), 2.09–2.23 that allometric effects should be considered when using pcl (female). Merus 0.92–1.13 pcl (male), 0.62–0.73 pcl walking leg morphometrics to separate P. stevensi from (female); length:height ratio 6.24–7.65 (male), 4.36–4.79 P. birsteini. The proportional lengths of the walking legs (female). Carpus 0.50–0.52 merus length (male), 0.57 of the smallest male P. stevensi (pcl 57.2 mm) approach (female). Propodus 0.79–0.87 merus length (male), those of the 79.5 mm male P. birsteini. 0.88–0.93 (female); length:height ratio 8.27–10.22 Unfortunately, most morphometric differences in (male), 6.55–7.06 (female). Dactylus 0.71–0.91 propodus the walking legs overlap between female P. stevensi and length (male), 0.75–0.89 (female). P. birsteini, so the most useful features distinguishing Pereopod 4 length 2.74–3.30 pcl (male), 2.12–2.17 females are those already mentioned above — differ- pcl (female). Merus 0.95–1.09 pcl (male), 0.61–0.62 pcl ences in dactyl length relative to the propodus, and (female); length:height ratio 6.60–7.86 (male), 4.70–5.00 differences in left and right cheliped height. (female). Carpus 0.46–0.52 merus length (male), 0.58– The smallest ovigerous P. stevensi and P. birsteini are 0.63 (female). Propodus 0.76–0.92 merus length (male), a similar size (pcl 62.4 mm and 61.1 mm, respectively), 0.93–0.99 (female); length:height ratio 8.45–11.22 and considerably larger than the smallest ovigerous (male), 6.72–7.52 (female). Dactylus 0.73–0.78 propodus P. gowlettholmes (pcl 28.8 mm). length (male), 0.76–0.91 (female). Distribution. Antarctica, from the Ross Sea and now Colour in life. Deep-red overall (Pl. 3G). the Amundsen Sea; 1389–1924 m.

Remarks. Ahyong & Dawson (2006) described Paralomis stevensi based on several damaged specimens from

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Paralomis taylorae sp. nov. reaching apex of article 4. Article 3 with 2 distal setose (Figs 105–108, Pl. 3F) tubercles. Scaphocerite a long, slender spine not overreaching distal peduncular article; with long lateral spine (one-third length of main spine), 3 shorter mesial Material examined. Holotype: TM G3609, female (cl spines and 1 or 2 scattered granules dorsally. Article 4 49.3 mm, pcl 39.8 mm, cw 37.5 mm), Tasman Sea, Hill unarmed, about half length of article 5. V, approx. 94 km SSE of Southeast Cape, Tasmania, Abdomen of female: Somite 2 covered with spines of 44°23’S, 147°09’E, 1310–1320 m, epibenthic sled, RV similar length to dorsal carapace spines. Somites 3–6 Southern Surveyor, SS01/97 stn 67, 31 Jan 1997. densely covered with short spines and acute granules, margins spinulate. Telson rounded, wider than long, Diagnosis. Carapace pyriform, slightly longer than with 6 blunt setose tubercles. wide, margins and surface densely covered with up- Pereopod 1 (chelipeds): Strongly spinose, unequal. right spines, surfaces glabrous; outer distance between Coxae unarmed, setose; distal margins with dense bases of anterolateral spines exceeding half carapace tufts of setae. Ischiobasis, merus, carpus, and propodus width; hepatic and branchial margins evenly merging, covered with arcuate spines and acute granules, longest without distinct constriction. Rostrum with 2 pairs of distally and dorsally. Dactylus and pollex with small dorsal spines; subrostral lobe absent. Distal dorsal eye- proximal spines, otherwise unarmed. spine extending anteriorly beyond cornea. Scaphocerite Major cheliped 1.78 pcl (female); upper palm length with 3 inner spines and 1 longer outer spine. Abdomen 1.59 times height (female); occlusal margins of fingers densely covered in short spines. Chelipeds and walking corneous for distal third, proximally with 3 calcare- legs covered with slender spines. Walking legs slender, ous nodules, proximal lowest; dactylus dorsal margin elongate, merus length exceeding 4 times height, about broadly convex, with tufts of golden setae and small 0.9 pcl; propodus length exceeding 9 times height proximal spine, 1.19 times longer than dorsal margin (exceeding 10 on walking leg 3); coxae unarmed or at of palm (female). most with a few blunt nodules. Minor cheliped 1.85 (female); upper palm length 1.75 times height (female); occlusal margin corneous Description. Carapace: Pyriform, 1.06 times longer than in quarter, proximally crenulate; dactylus dorsal mar- wide; regions distinct; surface and margins densely gin broadly convex, with tufts of golden setae and 5 covered with short, slender spines of varying length, small proximal spines, 1.75 times longer than dorsal and scattered granules; surface of spines glabrous; margin of palm. longest spine (on hepatic margin) 0.10 pcl; cervical Pereopods 2–4 (walking legs 1–3): Similar, elongate, groove distinct. Pterygostomian region sparsely spinu- spinose. Pereopod 2 longest. Coxae, unarmed, distal lose, spines widely spaced; with prominent, anterior, margins crenulate. Ischiobasis spinose. Merus ovate submarginal spine. in cross section; all surfaces spinose, spines longest Rostrum 0.24 pcl; broad basally, not markedly con- on extensor margins, shortest ventrally. Carpus and stricted proximal to dorsal spines; median spine slen- propodus spinose on all surfaces. Dactylus curved, der, ventral proximal margin with 4 minute spinules, laterally compressed; longer than carpus; proximal subrostral lobe absent; dorsal margin with 2 pairs of two-thirds spinose; flexor margin lined with 15–17 laterally divergent spines proximally, directed oblique- obliquely inclined corneous spinules; apex corneous. ly upwards; with scattered spinules and single short Pereopod 2 length 2.68 pcl (female). Merus 0.90 pcl spine between distal pair of dorsal spines. Posterior (female); length:height ratio 4.87 (female). Carpus 0.54 orbital margin concave, unarmed; outer orbital spine merus length (female). Propodus 0.88 merus length slender, directed anteriorly, not reaching beyond apex (female); length:height ratio 9.31 (female). Dactylus of cornea (when eyes directed anteriorly). Anterolateral 0.71 (female). spine shorter than outer orbital spine; outer distance Pereopod 3 length 2.80 pcl Merus 0.92 pcl (fe- between bases of anterolateral spines exceeding half male); length:height ratio 4.89 (female). Carpus 0.54 carapace width. merus length (female). Propodus 0.90 merus length Ocular peduncle: Longer than cornea; dorsally with (female); length:height ratio 10.15 (female). Dactylus scattered spines, distal spine longest, about half corneal 0.71 (female). diameter, overreaching cornea. Pereopod 4 length 2.74 pcl (female). Merus 0.85 Antennule: Peduncle unarmed, reaching anteriorly pcl (female); length:height ratio 4.35 (female). Car- beyond apex of antennal peduncle by quarter length pus 0.57 merus length. Propodus 0.99 merus length of distal antennular peduncle article. (female); length:height ratio 10.59 (female). Dactylus Antenna: Basal antennal article with curved ante- 0.70 (female). rolateral spine and distal spinule. Article 2 dorsally and laterally spinulose, with long distalolateral spine Colour in life. Bright red (Pl. 3F).

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 105. Paralomis taylorae sp. nov., female holotype, cl 49.3 mm, pcl 39.8 mm, cw 37.5 mm, Hill V, SSE of Southeast Cape, Tasmania (TM G3609). A, dorsal habitus. B, carapace. C, right pereopod 4.

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 106. Paralomis taylorae sp. nov., female holotype, cl 49.3 mm, pcl 39.8 mm, cw 37.5 mm, Hill V, SSE of South- east Cape, Tasmania (TM G3609). A, right chela. B, left chela. C, abdomen. D, posterior carapace and abdominal somite 2. E, carapace, right lateral view. F, anterior carapace, dorsal view.

Etymology. Named for Joanne Taylor, Museum Victo- Remarks. On the basis of its densely spinose dorsum, ria, for her usual helpfulness and assistance with the Paralomis taylorae sp. nov. resembles P. sonne Guzmán, Museum Victoria collections. 2009 [type locality: Chile], P. phrixa Macpherson, 1992 [type locality: Peru], P. poorei sp. nov., P. hystrixoides

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ 3 propodus length exceeding 10 times times height versus about 7 in P. sonne), in carapace proportions (slightly longer than wide rather than wider than long) and in having a sparsely, rather than a strongly and rather evenly spinose pterygostomial flap. Paralomis taylorae further differs from P. phrixa and P. hystrixoides in having unarmed or, at most, bluntly nodular distoventral margins of the pereopodal coxae, versus spinose in the latter two species. Paralomis taylorae further differs from P. spinosissima in its more slender walking legs in which the merus length exceeds 4 times its height and the propodus exceeds 9 times its height in contrast to 3–4 and 4–6 times height, respectively. The holotype of P. taylorae is a mature female, with setose ventral surfaces of the pereopodal coxae and a broad, well-developed abdomen.

Distribution. Presently known only from the type locality, off Southeast Cape, Tasmania; 1310–1320 m.

Paralomis webberi sp. nov. (Figs 81, 109–112)

Type material. Holotype: NMNZ Cr11141, ovigerous female (pcl 33.6 mm, cw 29.2 mm), Rumble III Seamount, Kermadec Ridge, 35°42.37’S, 178°28.69’E. 632–1255 m, trawled, 6 Sep 1998.

Figure 107. Paralomis taylorae sp. nov., female holotype, Diagnosis. Carapace pyriform, slightly longer than cl 49.3 mm, pcl 39.8 mm, cw 37.5 mm, Hill V, SSE wide; surface and margins densely and uniformly of Southeast Cape, Tasmania (TM G3609). A, right covered with slender spines of similar length; outer antenna, dorsal view. B, right antenna, ventral view. distance between bases of anterolateral spines slightly Scale = 5.0 mm. less than half carapace width. Subrostral lobe spinulate. Ocular peduncle dorsally spinulate, overreaching Sakai, 1980 [type locality: Japan], and P. spinosissima cornea. Scaphocerite with 3 long lateral spines and 3 Birstein & Vinogradov, 1972 [type locality: off South dorsal spines with longest about one-quarter length Georgia Islands, southwestern Atlantic]. Along with P. of longest lateral spine. Abdominal somites densely poorei sp. nov., the new species differs from the remain- covered with slender spines, becoming shorter on ing three species by lacking the prominent constriction successive somites. Chelipeds short, stout, outer and in the carapace margin at the junction of the hepatic upper surfaces densely spinose; mesial surface of palm and branchial regions, and in the proportionally wider with low, scattered, setose tubercles, otherwise smooth; anterior carapace margin, with the distance between dactyli proximally spinose. Walking legs similar, short, the bases of the anterolateral spines greater than half stout, not exceeding 1.5 pcl; segments densely spinose the maximum carapace width, rather than less than except for ventral surface of meri and coxae; pereopod half. Paralomis taylorae is readily distinguished from 4 merus length less than 2.5 times height; propodus P. poorei by its longer walking legs (pereopod 4 length length about 4.4 times height; dactylus shorter than in females exceeds 2.7 pcl versus less than 1.8 pcl) carpus, surface spinose for proximal two-thirds. Somite and more densely spinose carapace and walking legs 6 length 1.32 times width. (compare Figs 92 and 105; Tables 4, 5). In overall morphology, P. taylorae most closely resembles P. sonne in Description. Carapace: Pyriform, 1.15 times longer than sharing similar carapace and antennal ornamentation, wide; regions distinct; surface and margins densely and elongate walking legs. In addition to the anterior and uniformly covered with slender spines of simi- width of the carapace and lack of a prominent carapace lar length; surface of spines with minute, scattered, contriction, P. taylorae differs from P. sonne in having well-spaced simple setae; longest spine (on anterior more slender propodi of the walking legs (pereopod branchial margin) 0.18 pcl; cardiac region with 27

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spines; cervical groove distinct. Pterygostomian region above cornea; longest spine overreaching cornea by with slender anterior spine; spinulose on posterior length of cornea. two-thirds; surface of anterior third smooth except for Antennule: Peduncle unarmed, reaching anteriorly scattered granules near dorsal margin and with cluster beyond antennal peduncle by length of distal anten- of 6 or 7 spinules anteriorly. nular peduncle article. Rostrum 0.22 pcl; broad basally, not constricted Antenna: Basal antennal article with slender, curved, proximal to dorsal spines; median spine slender, sub- anterolateral spine, not reaching base of scaphocerite. rostral lobe angular, armed with 9 spinules; dorsally Article 2 with 2 inner distal spines; outer margin with with 2 median spines and pair of laterally divergent 3 or 4 slender spines and long distal spine reaching spines directed obliquely upwards. Posterior orbital distal quarter of ultimate peduncular article. Article margin concave, unarmed; outer orbital spine slender, 3 with small inner distal tooth. Scaphocerite a long, directed anterolaterally, reaching midlength of cornea. slender spine overreaching cornea spines; lateral Anterolateral spine as long as outer orbital spine; outer margins with 3 long spines; dorsally with 3 spines of distance between bases of anterolateral spines slightly about one-quarter length of longest lateral spine; inner less than half carapace width. margin unarmed. Article 4 unarmed, about one-third Ocular peduncle: Longer than cornea; with scattered length of article 5. granules and 10–12 dorsal spines, anterior 4 or 5 the Abdomen of female: Somites densely covered with longest, anteriorly directed, arranged in arcuate row slender spines, becoming shorter on successive somites.

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 109. Paralomis webberi sp. nov., ovigerous female holotype, pcl 33.6 mm, cw 29.2 mm, Rumble III Seamount (NMNZ Cr11141). A, dorsal habitus. B, carapace, right lateral view.

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 110. Paralomis webberi sp. nov., ovigerous female holotype, pcl 33.6 mm, cw 29.2 mm, Rumble III Seamount (NMNZ Cr11141). A, right chela outer surface. B, left chela, outer surface. C, left chela, inner surface. D, right pereopod 4, dorsal view. E, thoracic sternum and pereopodal coxa. F, right peropod 4, ventral view.

Spines on somite 2 similar to dorsal carapace spines. Pereopod 1 (chelipeds): Densely spinose, unequal. Spines of somites 3–6 less than half length of dorsal Major cheliped 1.31 pcl, 1.41 times height of minor carapace spines. Somite 6 length 1.32 times width. cheliped; minor cheliped 1.26 pcl. Coxae unarmed, Telson semicircular, with 8 blunt, apically setose tu- setose; distal margins with dense tufts of setae. Is- bercles. chiobasis spinose laterally, ventrally with clusters of

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 111. Paralomis webberi sp. nov., ovigerous female holotype, pcl 33.6 mm, cw 29.2 mm, Rumble III Seamount (NMNZ Cr11141). A, posterior carapace and abdominal somite 2. B, abdomen. C, anterior carapace, right lateral view. D, posterior abdominal somites and telson. E, carapace, anterior view. F, carapace, anteroventral view.

golden setae. Merus with smooth mesial surface, other for mesial surface with low scattered tubercles bearing surfaces spinose. Carpal surfaces densely spinose ex- tufts of golden setae. Fingers with tufts of golden setae; cept for mesial surface with clusters of golden setae. dactyli with proximal cluster of 5 spines. Fingers of Upper palm length 1.04 (major chela) and 1.11 times major chela with occlusal margins corneous for distal height (minor chela), surfaces densely spinose except quarter, proximally with 4 low calcareous prominences;

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 112. Paralomis webberi sp. nov., ovigerous female holotype, pcl 33.6 mm, cw 29.2 mm, Rumble III Seamount (NMNZ Cr11141). A, right antenna, ventral view. B, right antenna, dorsal view. C, rostrum, right lateral view. D, right eye, lateral view. E, right pereopod 4 dactylus. Scale = 1.0 mm.

dactylus 1.28 times dorsal margin of palm. Fingers of 1.00 merus length; length:height ratio 4.37. Dactylus minor chela with occlusal margins corneous for distal 0.65 propodus length. third to half, proximally crenulate; dactylus 1.64 times Colour in life. Not known. upper margin of palm. Pereopods 2–4 (walking legs 1–3): Similar, short, stout, Etymology. Named for Rick Webber, Museum of densely spinose. Pereopod 2 longest, 1.33 pcl. Coxae New Zealand Te Papa Tongarewa, for his helpfulness unarmed, surface setose, distal margins crenulate. Is- throughout the course of this work and for making the chiobasis spinose. Merus ovate in cross section, shorter large Te Papa lithodid collection available for study. than half pcl; dorsal, extensor and flexor surfaces spinose, longest extensor spines near distal third; ven- Remarks. Paralomis webberi sp. nov. closely resembles tral surface unarmed. Carpus spinose, longest spines P. histrix (de Haan, 1844) from Japan and P. echidna sp. as high as carpus. Propodus dorsoventrally flattened; nov. from southeastern Australia in the short walk- spinose, longest spines about 1.5 times propodal height. ing legs and densely spinose body and pereopods. Dactylus curved; shorter than carpus; surface spinose As with P. echidna, P. webberi is distinguished from for proximal two-thirds; distal third with tufts of P. histrix by the pattern of carapace spination. Although golden setae; flexor margin lined with 7 or 8 obliquely spine length varies between the mid-dorsal surface inclined corneous spinules; apex corneous. and carapace margins, for example, adjacent spines Pereopod 2 length 1.33 pcl. Merus 0.42 pcl; length: are generally of uniform or near uniform length in height ratio 2.62. Carpus 0.77 merus length. Propodus P. webberi and P. echidna, whereas in P. histrix, the 0.89 merus length; length:height ratio 4.37. Dactylus longer spines are interspersed with numerous shorter 0.65 propodus length. spines of about half the length of the former. Moreover, Pereopod 3 length 1.26 pcl. Merus 0.39 pcl; length: these shorter spines in P. histrix have rounded, rather height ratio 2.59. Carpus 0.80 merus length. Propodus than acute apices, and are often curved and inclined 0.91 merus length; length:height ratio 4.25. Dactylus anteriorly. All dorsal carapace spines of P. histrix are 0.66 propodus length. glabrous, rather than minutely setose as in P. webberi Pereopod 4 length 1.23 pcl. Merus 0.36 pcl; length: and P. echidna. Distinctions between P. webberi and height ratio 2.47. Carpus 0.84 merus length. Propodus P. echidna are subtle, as follows:

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ – The density of surface spination is greater in 668 m; but see Remarks below]. — Macpherson, 1988b: P. webberi than in P. echidna (at least in females). For 87–88, fig. 39, pl. 19B. — Zaklan, 2002: 773, 794. [New example, in P. webberi the cardiac region bears 27 synonymy]. spines versus 18–22 in P. echidna (in size-matched Acantholithus hystrix. — Dawson & Yaldwyn, 1985: 70 [misspelling; not A. histrix (de Haan, 1844)]. females). The differences are best observed, how- Paralomis zelandica. — McLay, 1988: 40–42, fig. 4. — Webber, ever, by direct comparison of specimens (Figs 74, 1997: 82. 109). Although spine length and density varies Paralomis hystrix. — McLay, 1988: 44. — Takeda in Amaoka allometrically in lithodids, the known females of et al., 1990: 362 [misspelling; not P. histrix (de Haan, P. webberi and P. echidna are of similar size, permit- 1844)]. ting direct comparison of size-related features. – The subrostral lobe is multispinulate in P. webberi, Type material. Holotype: NIWA 591, male (cl 111.6 mm, unarmed in P. echidna. pcl 97.0 mm, cw 101.6 mm), Chatham Rise, 44°18’S, – Abdominal somite 6 is more slender in P. webberi 174°31’E, 640 m, fine sandy mud, medium Agassiz than in P. echidna (length 1.32 times width versus trawl, Slope Benthos Cruise, FV Taranui, E0423, 15 1.08–1.15). Oct 1965. – The propodi of the walking legs are more slender in Other material examined. Holotype of Paralomis shinkai- P. webberi (length:height ratio 4.37 versus 3.68–3.73 maruae Takeda & Hatanaka, 1984: NSMT Cr8945, on pereopod 4 in females). Unfortunately, male female (pcl 75.4 mm, cw 78.1 mm), Bromley Plateau, P. webberi are not yet known. 31°13’30”S, 34°49’W, 668 m, RV Shinkai-Maru, 25 Mar Paralomis webberi superficially resembles juvenile 1977. P. zealandica in the dense dorsal spination. The most Cook Strait: NMNZ Cr11139, 1 male (cl 27.3 mm, pcl obvious distinguishing feature is size at maturity — 21.5 mm, cw 21.9 mm), S of Cape Turakirae, VUZ96, P. webberi is mature by at least 33.6 mm pcl whereas 28 Aug 1957; NMNZ Cr11140, 1 male (cl 27.9 mm, pcl P. zealandica matures at not less than about 60 mm 23.1 mm, cw 24.4 mm), off Cape Turakirae, VUZ111, pcl. Paralomis webberi is also readily distinguished 28 Jan 1958; NMNZ Cr11142, 2 males (cl 24.9–32.9 mm, from P. zealandica by its multispinulate subrostral pcl 19.0–25.7 mm, cw 18.3–24.5 mm), 1 female (cl 23.7 lobe (unarmed or unispinose in P. zealandica), more mm, pcl 18.9 mm, cw 18.5 mm), off Cape Turakirae, dense dorsal spination, absence of short spines on 41°30.2’S, 174°52’E, 380 m, BS424, RV Acheron, 11 Dec the inner surfaces of the palms of the chelipeds, and 1974. presence of spines on the proximal two-thirds rather Chatham Rise: NMNZ, 1 male (cl 120.9 mm, pcl 112.7 than proximal quarter to third of the extensor margins mm, cw 119.9 mm), Independent Island, 42°59.2’S– of the walking leg dactyli. Males of P. webberi are not 43°09.6’S, 174°41.1–07.4’E, 693–635 m, Tow 48, trip presently known, though they can be expected to have 1591, N. Mitchell, 9 Jan 2002; proportionally longer pereopods than in females, as in NIWA 42891, 1 female (pcl 79.4 mm, cw 78.9 other lithodids. mm), 43°07.29–05.71’S, 174°07.41–10.90’E, 668–690 Unfortunately, the specific habitat on Rumble III m, TAN0701/94, 15 Jan 2007; NMNZ Cr11750, 1 seamount from which the holotype of P. webberi was male (cl 112.9 mm, pcl 100.3 mm, cw 107.4 mm), 3 collected is not known. Rumble III is an active hy- ovigerous females (pcl 74.9–84.2 mm, cw 73.7–83.4 drothermal vent site, so P. webberi is possibly a vent mm), Mernoo Saddle, 43°14.42’S, 173°55.63’E, associate. 940–982 m, WIL/04/89, FV Will Watch, 23 Sep 1989; NIWA 42892,1 female (pcl 93.3 mm, cw 92.3 mm), Distribution. Presently known only from Rumble III 43°26.07–28.51’S, 174°14.52–16.90’E, 563–557 m, Seamount, southern Kermadec Ridge; 532–1255 m. TAN0701/96, 15 Jan 2007; NIWA 61193, 1 female (cl 110.5 mm, pcl 93.7 mm, cw 95.6 mm), 43°26.68’S, Paralomis zealandica Dawson & Yaldwyn, 174°05.50–05.82’E, 709–739 m, TAN0101/120, 21 Jan 1971 2001; NIWA 34927, 1 male (cl 121.0 mm, pcl 99.7 (Figs 113–119, Pl. 2F, 4F) mm, cw 99.7 mm), 43°51.7–57.4’S, 179°25.38–12.50’E, 495–574 m, trip 2413/89, coll. J. Williamson, 13 May Paralomis zealandica Dawson & Yaldwyn, 1971: 51–54 [type 2007; NMNZ Cr12022, 1 male (cl 87.3 mm, pcl 73.6 locality: Chatham Rise, 44°18’S, 174°31’E, 640 m]. — Daw- son & Yaldwyn, 1985: 70. — Dawson, 1989: 318. — O’Shea mm, cw 78.5 mm), 43°52.6–51.2’S, 179°14.4–17.4’W, et al., 1999: 49, fig. 17. — Zaklan, 2002: 774, 796. — Batson, 254–305 m, bottom temp. 4.5°C, FV Azuchi Maru, coll. 2003: 137. — Webber & Naylor, 2004b: 62. — Naylor et M. Yates, 12 Oct 1987; NMNZ, 1 male (cl 137.8 mm, al., 2005: 45–42. — Ahyong et al., 2007: 157. — Dawson, pcl 91.1 mm, cw 93.9 mm), 43°57.99’S to 44°00.00’S, 2008: fig. 7. 178°49.36–52.48’E, 638–675 m, TAN9212/074, 11 Paralomis shinkaimaruae Takeda & Hatanaka, 1984: 14–17, fig. Jan 1993; NIWA 3921, 2 males (cl 46.2–58.8 mm, pcl 3–5 [type locality: Bromley Plateau, 31°13’30”S, 34°49’W, 34.0–45.7 mm, cw 34.7–43.5 mm), 44°03.57–03.76’S,

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ 179°20.82–22.89’E, 677–692 m, TAN9812/95, Z9384, 27 FV Peterson, coll. D. Paterson & M. Miranovich, 11 Nov Oct 1998; NIWA 61192, 1 male (cl 101.7 mm, pcl 81.5 1994; NIWA 60564, 1 spent female (cl 87.0 mm, pcl 69.4 mm, cw 86.1 mm), 44°03.57–03.76’S, 179°20.82–22.89’E, mm, cw 70.3 mm), 51°12.33’S, 168°50.42’E, 552–555 m, 677–692 m, TAN9812/95, Z9384, 27 Oct 1998; NIWA, TAN0911/44, 8 Dec 2009; NIWA 60572, 1 male (cl 113.3 2 females (cl 106.4–118.7 mm, pcl 90.6–101.4 mm, cw mm, pcl 89.5 mm, cw 91.9 mm), 51°29.10’S, 173°06.03’E, 90–103.4 mm; larger ovigerous), 44°03.57–03.85’S, 544–545 m, TAN0911/25, 3 Dec 2009; NIWA 41572, 1 179°21.99–23.67’E, 673–688 m, TAN9812/94, Z9385, male (pcl 52.5 mm, cw 52.5 mm), 1 ovigerous female 27 Oct 1998; NIWA 61194, 1 female (pcl 87.8 mm, cw (pcl 90.1 mm, cw 95.5 mm), 1 damaged female (pcl 84.1 mm), 44°09.52–10.84’S, 179°55.36–51.57’W, 627–636 ~ 65 mm), N of Campbell Island, 51°44.04–43.00’S, m, TAN0101/67, 10 Jan 2001; NMNZ, 2 males (cl 75.9 168°35.61–40.17’E, 540–517 m, TAN0317/33, 26 Nov mm, pcl 60.1 mm, cw 63.1 mm; damaged, approx. pcl 2003; NMNZ Cr12023, 1 ovigerous female (pcl 112.9 80 mm), 44°12.9’S, 179°39.30’W–178°24.73’W, 598–560 mm, cw 115.0 mm), NE of Campbell Island, 52°18.75’S, m, TAN9105/26, 1 Jan 1992; NMNZ Cr5045, 1 male (cl 171°26.10’E, 497–565 m, trip 1994, tow 21, coll. R. 117.3 mm, pcl 103.2 mm, cw 106.8 mm) W of Chatham Hansen, 3 Oct 2004; NMNZ Cr11143, 1 female (cl Islands, 44°13.4–14.5’S, 178°14.4’W–177°53.1’W, 517 m, 71.2 mm, pcl 54.2 mm, cw 60.3 mm), Campbell Rise, FV Azuchi Maru, coll. M. Yates, 21 Oct 1987; NMNZ 53°00’S, 173°13’E, 508–510 m, RV Shinkai Maru, stn 269, Cr5897, 2 males (cl 118.5–120.8 mm, pcl 99.0–101.5 mm, 20 Jan 1977; NMNZ, 2 males (cl 94.3–116.3 mm, pcl cw 104.1–110.0 mm), 44°19.6–19.9’S, 173°36.6–37.0’E, 78.2–97.2 mm, cw 79.5–102.3 mm), SSW of Campbell 614–617 m, RV James Cook, J17/003/84, 29 Aug 1984; Island, 53°05.29–02.46’S, 168°55.36–53.95’E, 619–641 m, NMNZ, 1 male (cl 137.8 mm, pcl 117.4 mm, cw 127.2 TAN9105/60, 27 Nov 1991; NMNZ Cr6333, 1 male (cl mm), 44°20.80–21.64’S, 177°47.74’W–176°16.29’W, +112 mm, pcl 96.3 mm, cw 106.2 mm), SE of Campbell 626–631 m, TAN9106/39, 4 Jan 1992; NIWA 41407, 1 Island, 53°14.4–08.3’S, 170°35.5–22.6’E, 400 m, FV Ry- male (pcl 51.6 mm, cw 50.7 mm), 44°28.0’S, 175°47.0’E, uyu Maru, coll. H. Kavale, 30 Aug 1987; NIWA 60571, 688 m, Trip 2617, FV Amaltal Atlantis, coll. S. Yeo- 1 male (pcl 88.6 mm, cw 92.2 mm), 1 female (cl 117.3 man, 4 May 2008; NMNZ Cr9240, 1 male (cl 50.2 mm, pcl 96.7 mm, cw 98.4 mm), 53°23.37’S, 170°46.91’S, mm, pcl 38.3 mm, cw 37.4 mm), 44°28.81–29.71’S, 523–566 m, TAN0911/31, 5 Dec 2009; NIWA 61190, 2 179°45.53–48.26’E, 1210–1212 m, TAN9104/55, 21 Oct females (pcl 87.9–92.5 mm, cw 90.1–90.5 mm), Camp- 1991; NMNZ Cr11775, 1 male (cl 107.1 mm, pcl 86.7 bell Rise, 53°28.39–31.09’S, 171°04.09–06.01’E, 558 m, mm, cw 89.4 mm), SE of Banks Peninsula, 44°31.00– TAN0012/23, Z10600, 1 Dec 2000; NIWA 60570, 1 male 36.83’S, 173°38.00–26.92’E, 761–764 m, RV James Cook, (cl 103.1 mm, pcl 88.3 mm, cw 88.9 mm), 53°37.16’S, J15/13/83, 26 Nov 1983. 170°44.94’S, 776–778 m, TAN0911/32, 5 Dec 2009; Bounty Plateau: NMNZ Cr11776, 1 female (cl 92.9 NMNZ Cr11768–11770, 3 females (cl 79.8–113.2 mm, mm, pcl 77.8 mm, cw 81.5 mm), SE of Bounty Islands, pcl.64.1–105.4 mm, cw 65.2–107.0 mm), Auckland to 48°15.5–19.0’S, 179°48.5’E–180°00.0’E, 512–503 m, RV Campbell Islands, Feb 1992; NSMT Cr10117, 1 male (cl Shinkai Maru, stn 79, coll. J.G. Jones, 17 Nov 1978; 42.9 mm, pcl 33.0 mm, cw 31.9 mm), Campbell Plateau, NMNZ Cr11719–11720, 2 males (cl 123.6–124+ mm, pcl 542 m, T-142, 8 Nov 1983. 106.9–114.9 mm, cw 112.3–124.6 mm), SW of Bounty New Zealand, no specific data: NMNZ Cr11575, 2 Islands, 48°01–05’S, 178°32–34’E, 545–454 m, RV Shinkai males (cl 105.0–118.6 mm, pcl 89.1–100.5 mm, cw Maru, 12/78 stn 239, 11 Jan 1979. 90.8–105.8 mm), New Zealand, no other data; NMNZ Campbell Plateau: NIWA 60575, 1 male (pcl 85.0 Cr11576, 1 male (cl 104.0 mm, pcl 88.2 mm, cw 95.5 mm, cw 89.0 mm), 46°59.35’S, 169°58.25’E, 729–710 mm), New Zealand. m, TAN0911/3, 27 Nov 2009; NIWA 61191, 1 male (pcl 87.9–92.5 mm, cw 90.1–90.5 mm), 1 female (pcl Diagnosis. Carapace pyriform, slightly longer to 87.9–92.5 mm, cw 90.1–90.5 mm), Campbell Plateau, slightly wider than long, margins and surface densely 49°12.82–12.22’S, 167°53.74–58.24’E, 702–685 m, covered with conical spines, surfaces glabrous, apices TAN0012/45, Z10601, 7 Dec 2000; NMNZ Cr5896, 1 with few setae in early adults, corneous in late adults; male (pcl 94.2 mm, cw 96.5 mm), 2 females (larger outer distance between bases of anterolateral spines ovigerous) (cl 98.7–114.7 mm, pcl 79.4–92.9 mm, cw distinctly less than half carapace width. Rostrum 77.7–94.0 mm), 49°19.2–17.8’S, 173°38.4–39.4’E, 597–617 prominently trispinous; subrostral lobe rounded or m, RV James Cook, J04/001/84, 17 Feb 1984; NMNZ produced to small spine. Scaphocerite prominently Cr4857, 1 male (cl 68+ mm, pcl 60.9 mm, cw 58.5 mm), multispinose. Abdomen densely covered with conical NW of Urry Bank, 49°30.6–39.54’S, 174°08.6–01.8’E, 798 spines. Pereopods covered with spines, most elongate m, FV Oyang, #7, Tow 5, coll. C.D. Roberts, 7 May 1987; on dorsal and extensor margins; dactylus longer than NMNZ Cr11146, 1 female (pcl 50.73 mm, cw 51.29 mm), carpus; spines on dactylar extensor margin restricted E of Auckland Islands, 50°50’S, 166°52’E, 390–400 m, to proximal one-third of margin.

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 113. Paralomis zealandica Dawson & Yaldwyn, 1971, male, pcl 94.2 mm, Campbell Plateau (NMNZ Cr5896). A, dorsal habitus. B, carapace, right lateral view.

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 114. Paralomis zealandica Dawson & Yaldwyn, 1971, male, pcl 94.2 mm, Campbell Plateau (NMNZ Cr5896). A, carapace. B, ventral surface and abdomen. C, anterior carapace, dorsal view. D, posterior carapace and abdominal somite 2.

Description. Carapace: Pyriform, 0.90–1.05 times longer proximally with 2 pairs of laterally divergent spines than wide; regions distinct; surface and margins uni- directed obliquely upwards and single upright spine formly covered with short, conical spines (longer and between proximal pair of dorsal spines. Posterior slender in early adults and juveniles); spines with orbital margin concave, unarmed; outer orbital spine smooth surface and several apical simple setae, which slender, directed anteriorly, overreaching eyes. Ante- are lost in largest specimens; longest spine (on anterior rolateral spine as long as or longer than outer orbital branchial margin) 0.07–0.26 pcl; cardiac region with spine; outer distance between bases of anterolateral 13–15 spines; cervical groove distinct. Pterygostomian spines distinctly less than half carapace width. region with acute granules and small spines; with Ocular peduncle: Longer than cornea; dorsally with prominent anterior spine. short, scattered granules or spines, distalmost spine Rostrum 0.07–0.31 pcl; broad basally, not con- longest; distal spine about half corneal diameter, over- stricted proximal to dorsal spines; median spine slen- reaching cornea. der, ventral lobe bluntly angular or with sharp apex;

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 115. Paralomis zealandica Dawson & Yaldwyn, 1971, male, pcl 94.2 mm, Campbell Plateau (NMNZ Cr5896). A, right chela. B, left chela. C, anterior carapace, right lateral view. D, left pereopod 4 dactylus. E, left pereopod 4.

Antennule: Peduncle unarmed, reaching anteriorly article; with 2 long lateral spines, first minute, second slightly beyond apex of antennal peduncle by three- about two-thirds length of main spine; dorsomesially quarters to full length of distal antennular peduncle with 2 spines, shorter than half length of lateral spines. article. Article 4 unarmed, about half length of article 5. Antenna: Basal antennal article with slender, arcuate Abdomen: Ornamentation of both sexes similar. anterolateral spine. Article 2 with slender inner spine, Somites 2–6 covered with spines of similar length to occasionally bifid; outer margin with 2–4 small spines dorsal carapace spines. Telson with 2 or 3 pairs of stout proximally and long, distal spine overreaching article 3. spines; semicircular. Article 3 unarmed. Scaphocerite multispinose; primary Pereopod 1 (chelipeds): Strongly spinose, unequal; spine long, slender, overreaching distal peduncular ornamentation similar on both sides and in both sexes.

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 116. Paralomis zealandica Dawson & Yaldwyn, 1971, ovigerous female, pcl 92.9 mm, Campbell Plateau (NMNZ Cr5896). A, dorsal habitus. B, left pereopod 4. C, carapace. D, ventral surface and abdomen.

Coxal margins spinose. Ischiobasis spinose laterally for distal third, proximally with 3 calcareous nodules; and ventrally. Merus with relatively smooth mesial dactylus dorsal margin broadly convex, with rows of surface, other surfaces spinose. Carpus prominently tufts of golden setae and proximal cluster of 3–6 small spinose, without setal clusters. Palm with most surfaces spines, 0.93–1.10 times longer than dorsal margin of prominently spinose; inner (mesial surface) sparsely palm (male), 1.14–1.24 (female). spinose, with some apically setose. Minor cheliped 1.30–1.37 pcl (male), 1.07–1.11 (fe- Major cheliped 1.51–1.68 pcl (male), 1.13–1.18 (female); upper palm length 1.04–1.14 times height (male), male); upper palm length 0.94–1.02 times height (male), 1.02–1.06 (female); occlusal margin corneous in distal 0.98–1.06 (female); occlusal margins of fingers corneous half, proximally crenulate; dactylus dorsal margin

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 117. Paralomis zealandica Dawson & Yaldwyn, 1971. A–D, male holotype, cl 111.6 mm, pcl 97.0 mm, cw 101.6 mm, Chatham Rise (NIWA 591). E, female, pcl 92.5 mm, Campbell Rise (NIWA 61190). F, female, pcl 87.9 mm, Campbell Rise (NIWA 61190). A, dorsal habitus. B, ventral surface and abdomen. C–D, left pereopod 4, dorsal and ventral views. E–F, anterior carapace, right lateral view.

broadly convex, with rows of tufts of golden setae and Pereopods 2–4 (walking legs 1–3): Similar, elongate, proximal cluster of 2–4 small spines, 1.49–1.57 times spinose. Pereopod 3 longest. Coxae distally spinose; longer than dorsal margin of palm (male), 1.52–1.61 ventral surfaces with short spines or acute tubercles, be- (female). coming low and blunt in mature females and obsolete

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Figure 118. Paralomis zealandica Dawson & Yaldwyn, 1971, A–B, dorsal and ventral views, female, pcl 54.2 mm, Campbell Rise (NMNZ Cr11143). C–D, dorsal and ventral views, male, pcl 25.7 mm, Cook Strait (NMNZ Cr11142). E–F, dorsal and right lateral views, male, pcl 21.5 mm, Cook Strait (NMNZ Cr11139).

on pereopod 4. Ischiobasis spinose. Merus subovate dorsoventrally flattened, shorter than merus; extensor in cross section; dorsal surface, extensor and flexor and flexor margins each with row of 6–8 spines; dorsal margins prominently spinose, each with relatively and ventral surfaces with 2 relatively even rows of 6–8 even rows of 7–12 spines; ventral surface with a row of spines. Dactylus curved, laterally compressed; longer small spines or acute tubercles near flexor and exten- than carpus (subequal in juvenile males < 30 mm pcl); sor margins. Carpus spinose on all surfaces. Propodus with cluster of spines on proximal quarter to third,

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ and scattered tufts of golden setae along length; flexor (male), 0.80–0.86 (female); length:height ratio 5.39–5.62 margin lined with 7–9 obliquely inclined corneous (male), 3.57–4.26 (female). Dactylus 0.96 propodus spinules; apex corneous. length (male), 0.81–0.90 (female). Pereopod 2 length 2.31–2.34 pcl (male), 1.49–1.66 Pereopod 3 length 2.44–2.53 pcl (male), 1.49–1.69 pcl (female). Merus 0.73–0.78 pcl (male), 0.47–0.51 pcl pcl (female). Merus 0.73–0.80 pcl (male), 0.45–0.53 pcl (female); length:height ratio 3.22–3.74 (male), 2.33–2.61 (female); length:height ratio 3.12–3.81 (male), 2.45–2.78 (female). Carpus 0.60–0.63 merus length (male), (female). Carpus 0.60–0.64 merus length (male), 0.67– 0.68–0.76 (female). Propodus 0.77–0.81 merus length 0.74 (female). Propodus 0.79–0.84 merus length (male),

Figure 119. Geographic and bathymetric distribution of Paralomis zealandica Dawson & Yaldwyn, 1971.

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ 0.81–0.87 (female); length:height ratio 5.32–6.01 (male), the proximal half. Features distinguishing P. zealandica 3.67–4.44 (female). Dactylus 0.92–0.97 propodus length from P. poorei, P. echidna, and P. webberi are given under (male), 0.78–0.93 (female). the respective accounts of the latter three species. Pereopod 4 length 2.31–2.42 pcl (male), 1.46–1.68 Other allometric changes in P. zealandica are in pcl (female). Merus 0.53–0.74 pcl (male), 0.35–0.50 pcl the proportional lengths of the walking legs, which (female); length:height ratio 2.98–4.01 (male), 2.48–2.81 are proportionally longest in large males, and in the (female). Carpus 0.63–0.72 merus length (male), 0.59– proportions of the walking leg meri, which are more 0.76 (female). Propodus 0.84–0.88 merus length (male), slender in the largest males (length:height ratio 2.7–2.8 0.89–0.92 (female); length:height ratio 5.69–6.22 (male), at 19.0–25.7 mm pcl, 3.5–4.0 at 91.1–103.2 mm pcl). The 3.76–4.60 (female). Dactylus 0.96–0.99 propodus length walking leg dactyli are about as long as the carpus (male), 0.83–0.90 (female). length in juveniles < 30 mm pcl, and longer than the carpus in larger specimens. The ventral surfaces of the Colour in life. Overall ivory-white, often with pale coxae of the chelipeds and walking legs are armed with red-orange mottling on pereopods. Carapace with acute tubercles in juveniles of both sexes and retained spines on gastric and cardiac regions, and usually also in adult males. In females, however, the ventral coxal branchial regions, red-orange. (Pl. 2F). tubercles become obsolete as the abdomen expands with maturity, covering the coxae. The smallest oviger- Remarks. Paralomis zealandica Dawson & Yaldwyn, ous female is 74.9 mm pcl, although specimens appear 1971, is the most widespread species of the genus in to be mature by about 60 mm pcl. New Zealand, ranging from the Chatham Rise to the Takeda & Hatanaka (1984) described P. shinkai- Bounty and Campbell plateaus. It attains a large size maruae from Bromley Plateau, southwestern At- and undergoes significant allometric changes in dorsal lantic, based on an ovigerous female collected by spination. Juveniles are densely covered in slender the RV Shinkai-Maru, apparently in March 1977. spines, superficially resembling other small, spiny Distinguishing P. shinkaimaruae from adult species such as the Japanese P. histrix (de Hann, 1844) P. zealandica, however, is problematical. The holo- (frequently misspelt as P. hystrix), P. echidna sp. nov. type of P. shinkaimaruae differs from the holotype of (southeastern Australia), P. webberi sp. nov. (Kermadec P. zealandica in the presence of a small subrostral Ridge, New Zealand), and P. poorei sp. nov. (southern spine rather than a blunt lobe. This feature, however, Australia and New Zealand). The dorsal spines of is variable in P. zealandica, with the subrostral lobe P. zealandica become proportionally shorter with in- often being produced into a small spine. Thus, the creasing body size, becoming short conical tubercles two species cannot be distinguished and are herein in adults. These allometric changes in spination of synonymised. The resulting disjunct distribution of P. zealandica have led to erroneous reports of the P. zealandica, however, is anomalous. It is not adequate- Japanese P. histrix from New Zealand (e.g. Dawson ly explained by sampling gaps because regular fishing & Yaldwyn 1985; Takeda in Amaoka et al. 1990). The effort in intermediate regions is likely to have yielded specimen on which the original New Zealand report of further specimens. Likewise, the prospect of artificial P. histrix was based (NMNZ Cr11143, Fig. 118A–B) is an introduction is unlikely. A more parsimonious explana- immature female of P. zealandica. Similarly, Takeda in tion is that the holotype of P. shinkaimaruae originated Amaoka et al. (1990) clearly depicts P. zealandica (as P. from New Zealand, probably the Bounty Plateau, hystrix). As suggested by McLay (1988), New Zealand where the Shinkai-Maru was operating in early 1977. records of P. histrix are based on juvenile P. zealandica. The specimen was probably mislabelled or mixed-up Paralomis zealandica and P. histrix superficially resem- with samples collected only one to two months later ble each other both as juveniles and adults because when the Shinkai-Maru continued operations on the both species undergo similar allometric reduction Bromley Plateau, southwestern Atlantic. in spination and attain a similar adult size. Paralomis Paralomis zealandica occurs only in southeastern zealandica, however, is readily distinguished from New Zealand waters, from Cook Strait to the Campbell P. histrix by its more uniform length carapace spines Plateau. (versus long spines interspersed with numerous, anteriorly recurved, apically rounded spines of about half Distribution. Southeastern New Zealand, from Cook the length of the former, Fig. 82), and in the extensor Strait to the Campbell Plateau; 254–1212 m, usually marginal spines of the dactyli of the walking legs being 500–700 m. restricted to the proximal quarter to third, rather than

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Prior to the present study, a total of 10 extant species the presumed circumpolar distribution of L. murrayi of king crab were believed to occur in New Zealand (see also Macpherson 1988b, c). At present, L. murrayi and Australia: Lithodes murrayi Henderson, 1888, is reliably known only from the southwestern Indian L. longispina Sakai, 1971, Neolithodes brodiei Dawson Ocean. Wide ranges within Lithodidae, at least in the & Yaldwyn, 1970, N. flindersi Ahyong, 2010, and six western Pacific, appear to be the exception rather the species of Paralomis. Seven lithodid species had been rule, and ranges of all putatively widespread lithodids reported from New Zealand waters, and six from should be examined closely. Australia (including Macquarie Island) of which three Many species of lithodid from the study area are were believed to occur on both sides of the Tasman known from few specimens, so little can generally Sea. Three species in two genera of lithodid have been be inferred about their distributions: L. chaddertoni, recorded from the Ross Sea, Antarctica. A total of 23 L. rachelae, P. webberi, and P. taylorae are each known species in three genera are now known from the study only from a single locality or localities in close geo- area: 13 species in New Zealand, 12 in Australia (four graphic proximity. Other species, such as P. echidna, shared by both regions) and three in the Ross Sea (wth P. poorei, and P. staplesi, also known from only a few Paralomis birsteini Macpherson, 1988a extending onto sites, have been collected from widely disparate lo- the Macquarie Ridge). The Macquarie Ridge record of calities on both sides of the Tasman Sea, and can be P. birsteini is significant in providing a biogeographic regarded as relatively wide ranging. Lithodes robertsoni link between Antarctic P. birsteini and P. stevensi, and is apparently relatively widespread in New Zealand, the very similar P. gowlettholmes sp. nov. from southern from the Challenger Plateau, Chatham Rise south to Tasmania. Results of the present study demonstrate the Campbell Plateau. Lithodes jessica is known from that rather than being relatively depauperate in com- the southern Lord Howe Rise and off Hawkes Bay, parison to the north Pacific, and presumed centre of New Zealand. Similarly, L. richeri and P. dawsoni have origin of the group, the lithodid fauna of the south- relatively wide southwestern Pacific ranges, from the western Pacific is considerably richer than previously Coral Sea south to southeastern Australia and New known. Zealand, respectively. This is probably also true for That only four lithodid species are shared by N. bronwynae if records of N. vinogradovi from New Australia and New Zealand (Lithodes macquariae sp. Caledonia prove referable to the former, as suggested nov., Paralomis echidna sp. nov., P. poorei sp. nov., above. Species for which the most substantial distri- P. staplesi sp. nov.) implies limited dispersal potential butional data exist are L. aotearoa, N. brodiei, and P. of the regional lithodids. Larvae of some lithodids, zealandica from New Zealand, and L. australiensis, N. such as Paralithodes camtschaticus, are planktotrophic flindersi, and P. gowlett-holmes from Australia. and therefore likely to disperse via ocean currents, but The ranges of the two most common and most wide- larvae of Lithodes, Paralomis, and probably Neolithodes, spread lithodids in New Zealand waters, L. aotearoa and are lecithotrophic, limiting prospects for pelagic disper- N. brodiei, follow the deep shelf and slope surrounding sal (Shirley & Zhou 1997; Thatje et al. 2005). Neolithodes mainland New Zealand, usually at depths of around brodiei and L. longispina were previously thought to oc- 1000 m, principally the southern part of the West Nor- cur on both sides of the Tasman Sea, but present results folk Ridge, the Challenger Plateau, Chatham Rise and indicate that Australian records of N. brodiei are refera- Campbell Plateau — essentially the ancient landmass of ble to N. flindersi, and records of L. longispina from New Zealandia. The two most common species of Paralomis Zealand and Australia are based on a different new spe- in New Zealand waters, P. dawsoni and P. zealandica, ap- cies in each region (L. aotearoa, L. australiensis). Lithodes parently have an additional oceanographic component longispina, itself, previously believed to be widespread affecting their distributions. Paralomis dawsoni occurs (Japan to Guam, New Zealand and Australia) (Dawson along undersea ridges extending from the Coral Sea 1989), has been shown to probably occur only in the (Solomon Islands and New Caledonia) south to the Japanese region, following recognition of L. aotearoa, Chatham Rise. Paralomis zealandica ranges from Cook L. robertsoni, and L. australiensis herein, in addition Strait and the Chatham Rise south to the Campbell to similar species in Guam (L. paulayi Macpherson & Plateau. The Chatham Rise acts as the intersection Chan, 2008) and Taiwan (L. formosae Ahyong & Chan, between the warm, south-flowing Tasman Front origi- 2010). Likewise, the discovery that previous records of nating in the Coral Sea, and the colder north-flowing L. murrayi from Australia and New Zealand are refer- subantarctic inflow. This intersection of water masses, able to L. aotearoa and L. macquariae further undermines the Subtropical Convergence, corresponds to the distri-

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ butional overlap of P. dawsoni and P. zealandica. More- and P. birsteini, which occur at depths approaching or over, this distributional ‘boundary’ between P. dawsoni exceeding 2000 m and have trans-continental ranges. and P. zealandica also corresponds to broader patterns The lithodid fauna of New Zealand and Australia of benthic diversity and may be regarded as a general is now known to be about twice as large as previous biogeographic boundary (Nodder et al. 2003). estimates. In the present study, emphasis on limb The ranges of the two most common Australian morphometrics has proven particularly useful in dis- lithodids, L. australiensis and N. flindersi, essentially tinguishing species when considered in the context of follow the outer shelf and continental slope off south- allometry and sexual dimorphism. Also, for Lithodes, eastern Australia. Lithodes australiensis also occurs the spination of the walking leg dactyli, spination of the along the southeastern coast from New South Wales posterior branchial margin, and fusion of the plates of south to eastern Tasmania and onto the South Tas- abdominal somite 2 have also proven to be taxonomi- man Rise. Neolithodes flindersi ranges from New South cally useful. These featutres should be considered in Wales around Tasmania and into the Great Australian any future studies of Lithodes. The large collections Bight. Paralomis gowlettholmes is presently known only available for study have permitted most species to be from the southern Tasmanian seamounts, and appears documented from both sexes over a wide size range. to have its nearest relatives in Antarctica and on the Some species, however, remain to be more adequately Macquarie Ridge (P. birsteini). characterised, such as Lithodes robertsoni and L. chad- Because most species of Neolithodes and Lithodes in dertoni, being known only from very few specimens. New Zealand and Australia do not occur beyond slope Males of Paralomis taylorae and webberi are not presently depths, their geographic distributions are related to the known, nor are females of Neolithodes bronwynae and extent of contiguous continental shelf and slope habitat. Paralomis staplesi. Clearly, further research into the Consequently, the Australian and New Zealand species regional lithodid fauna is still required. It is also likely of Neolithodes and Lithodes are isolated and endemic to that more species will be discovered from the region, their respective regions (though it is conceivable that especially in northern New Zealand waters and south- some southern overlap might occasionally occur via the western Australia. Northern Australian waters remain South Tasman Rise and Macquarie Ridge, and northern unstudied for Lithodidae. communication might occur via the various ridge sys- tems in the northern Tasman Sea). In contrast, species that occur down to abyssal depths can be expected to range more widely, as observed in Paralomis staplesi

ACKNOWLEDGMENTS

This work was largely supported by the Founda- ecosystems (C01X0224), Biodiversity of the western tion for Research Science and Technology (contract Ross Sea (ZBD200303), IPY-CAML project (LIN08302), C01X0502) and the NIWA Capability Fund. I wish to Oceans Survey 2020, and various trawl surveys (project thank Rick Webber (NMNZ), Stephen Keable (AM), MDT2007-01C). Malcom Clark, Karen Gowlett- Joanne Taylor and Gary Poore (NMV), Tin-Yam Chan Holmes, Sadie Mills, Reyn Naylor, Steve O’Shea, (NTOU), Thierry Laperousaz (SAM), Roger Buttermore Ashley Rowden, Kareen Schnabel, Darren Stevens, (TM), Peter Ng (ZRC), Hironori Komatsu (NSMT), Dean Stotter, and Di Tracey are thanked for their help Rafael Lemaitre (USNM), and Régis Cleva (MNHN) in numerous ways, including provision of colour im- for making their collections available for study. Many ages of selected species. Enrique Macpherson, Colin of the specimens studied here were collected under McLay, and Pat Mclaughlin are also thanked for valu- numerous research programmes funded by the New able comments and useful discussions, as are Dennis Zealand Ministry of Fisheries and Foundation for Re- Gordon and Tracy Farr for their careful copy-editing. search Science and Technology, including the following Lastly, special thanks to Rachel for her tireless sup- programmes: Ocean Ecosystems (C01X0223, C01X027), port. Seamounts: their importance for fisheries and marine

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Principal taxonomic account is in bold font; species illustrations in bold italic. Synonymic entries and material examined are not included. Acantholithus 109 Lithodes maja 15 amphipods 116 Lithodes mamillifer 15, 16, 55 Alvinocaris alexander 148 Lithodes manningi 15 Alvinocaris longirostris 148 Lithodes megacantha 15, 16, 48 Alvinocaris niwa 148 Lithodes murrayi 6, 26, 27, 54, 55, 116, 185 Anomura 5, 9 Lithodes nintokuae 15 Lithodes panamensis 15 Brachyura 5 Lithodes paulayi 15, 16, 37, 48, 60, 70, 72, 185 Briarosaccus callosus (parasitic rhizocephalan) 115, 147, Lithodes rachelae 16, 36, 37, 41, 48, 55, 56, 57, 59, 70, 97, 148 185 Lithodes richeri 16, 36, 47, 60, 61, 63, 65, 97, 185 Careproctus sp. (liparid fish) 83, 96 Lithodes robertsoni 16, 36, 37, 48, 54, 60, 66, 67, 68, 69, Chrysogorgia orientalis (coral) 96 71, 72, 97, 185, 186 Cryptolithodes 14 Lithodes santolla 5, 8, 16 Lithodes turkayi 16, 55 Echinocerus 109 Lithodes turritus 16 Lithodes unicornis 16 Gandalfus puia 148 Lithodes wiracocha 16 Glyptolithodes 15 Lopholithodes 15, 109

Hapalogastridae 5, 9, 14 Nautilocaris saintlaurentae 148 Neolithodes 8, 9, 10, 11, 13, 15, 73 isopods 116 Neolithodes agassizi 73 Neolithodes asperrimus 73 Lebbeus wera 148 Neolithodes brodiei 6, 74, 75, 76, 77, 79, 81, 89, 96, 98, Leptolithodes 109 185 Lithodoidea 14 Neolithodes bronwynae 74, 82, 83, 84, 85, 86, 88, 98, 185, Lithodidae 14 186 Lithodes 8, 10, 11, 13, 14, 15, 186 Neolithodes capensis 73, 107 Lithodes aequispinus 5, 15 Neolithodes diomedeae 73 Lithodes aotearoa 9, 15, 16, 17, 19, 22, 23, 25, 27, 36, 37, Neolithodes duhameli 73, 87, 89 41, 48, 55, 72, 73, 83, 97, 100, 148, 185 Neolithodes flindersi 6, 74, 83, 89, 90, 91, 92, 93, 95, 98, Lithodes australiensis 15, 16, 29, 30, 31, 33, 34, 41, 48, 55, 101, 185, 186 60, 64, 70, 72, 97, 185, 186 Neolithodes grimaldii 73 Lithodes ceramensis 15, 16 Neolithodes nipponensis 73, 83, 96 Lithodes chaddertoni 13, 15, 16, 36, 37, 38, 39, 41, 48, 100, Neolithodes vinogradovi 73, 87, 89, 185 185, 186 Neolithodes yaldwyni 6, 74, 83, 98, 100, 101, 103, 104, Lithodes confundens 8, 15 105 Lithodes ferox 15 Lithodes formosae 15, 16, 185 Paguroidea 5 Lithodes galapagensis 15, 16 Paralithodes 15 Lithodes granulosa 109 Paralithodes camtschaticus 5, 185 Lithodes histrix 109 Paralomis 8, 9, 10, 11, 13, 15, 108, 185 Lithodes jessica 13, 16, 40, 41, 42, 43, 44, 45, 46, 185 Paralomis aculeata 108, 109 Lithodes longispina 6, 15, 26, 27, 36, 64, 185 Paralomis africana 108 Lithodes aff. longispina 6, 60 Paralomis alcockiana 108, 124 Lithodes macquariae 15, 16, 48, 49, 50, 51, 53, 98, 155, Paralomis anamerae 108 185 Paralomis arae 108

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This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-nd/3.0/ Paralomis arethusa 108 Paralomis manningi 108 Paralomis aspera 108 Paralomis medipacifica1 08 Paralomis birsteini 6, 99, 100, 108, 109, 110, 111, 112, 113, Paralomis mendagnai 108 140, 141, 155, 158, 159, 165, 185, 186 Paralomis microps 108 Paralomis cf. birsteini 6 Paralomis multispina 108, 109 Paralomis bouvieri 108, 109, 154 Paralomis nivosa 108 Paralomis ceres 108 Paralomis ochthodes 109 Paralomis chilensis 108 Paralomis odawarai 109 Paralomis cristata 108 Paralomis otsuae 109 Paralomis cristulata 108, 109 Paralomis pacifica1 09 Paralomis cubensis 108, 124 Paralomis papillata 109 Paralomis danida 108 Paralomis pectinata 109 Paralomis dawsoni 6, 8, 99, 108, 109, 110, 116, 117, 118, Paralomis phrixa 109, 168, 169 119, 120, 122, 123, 185, 186 Paralomis cf. phrixa 6 Paralomis debodeorum 6 Paralomis poorei 99, 109, 110, 133, 134, 148, 149, 150, 151, Paralomis diomedeae 108 152, 153, 154, 160, 168, 169, 184, 185 Paralomis dolfleini1 08, 109 Paralomis roeleveldae 109 Paralomis echidna 108, 109, 110, 125, 126, 127, 128, 129, Paralomis seagranti 109, 123, 124 130, 131, 132, 155, 160, 174, 175, 184, 185 Paralomis serrata 109 Paralomis elongata 108 Paralomis shinkaimaruae 109, 184 Paralomis erinacea 108 Paralomis sonne 109, 168, 169 Paralomis formosa 108, 109 Paralomis spectabilis 6, 109, 115, 140 Paralomis gowlettholmes 99, 108, 109, 110, 115, 116, 135, Paralomis spinosissima 109, 169 136, 137, 138, 139, 155, 158, 159, 165, 185, 186 Paralomis staplesi 99, 109, 110, 116, 133, 155, 156, 157, Paralomis granulosa 108, 109 158, 159, 185, 186 Paralomis grossmani 108 Paralomis stella 109 Paralomis haigae 108 Paralomis stevensi 6, 99, 109, 110, 115, 116, 140, 141, 155, Paralomis heterotuberculata 109 158, 159, 160, 161, 162, 163, 164, 185 Paralomis hirtella 6, 8, 100, 108, 110, 133, 142, 143, 144, Paralomis taylorae 99, 109, 155, 166, 167, 168, 169, 170, 145, 146 185, 186 Paralomis histrix 6, 108, 109, 123, 133, 134, 174, 174, Paralomis truncatispinosa 109 184 Paralomis tuberipes 109 Paralomis cf. histrix 6, 134 Paralomis verrilli 109 Paralomis hystrixoides 108, 168, 169 Paralomis webberi 109, 110, 133, 155, 169, 171, 172, 173, Paralomis inca 108 174, 184, 185, 186 Paralomis indica 108 Paralomis zealandica 6, 9, 98, 100, 109, 110, 123, 125, 155, Paralomis investigatoris 108 175, 176, 178, 179, 180, 181, 182, 185, 186 Paralomis japonica 108 Pristopus 109 Paralomis jamsteci 108 Pristopus verrilli 109 Paralomis aff. jamsteci 147 Paralomis kyushupalauensis 108 Rhinolithodes 15 Paralomis longidactylus 108 Paralomis longipes 108, 109 Sculptolithodes 15 Paralomis makarovi 108, 154

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